Transitions in functional morphology from “large branchiopods” to Cladocera: Video and confocal microscopic studies of Cyclestheria hislopi (Cyclestherida) and Sida crystallina (Cladocera: Ctenopoda)

Great diversity is found in morphology and functionality of arthropod appendages, both along the body axis of individual animals and between different life-cycle stages. Despite many branchiopod crustaceans being well known for displaying a relatively simple arrangement of many serially post-maxillary appendages (trunk limbs), this taxon also shows an often unappreciated large variation in appendage morphology. Diplostracan branchiopods exhibit generally a division of labor into locomotory antennae and feeding/filtratory post-maxillary appendages (trunk limbs). We here study the functionality and morphology of the swimming antennae and feeding appendages in clam shrimps and cladocerans and analyze the findings in an evolutionary context (e.g., possible progenetic origin of Cladocera). We focus on Cyclestheria hislopi (Cyclestherida), sister species to Cladocera and exhibiting many “large” branchiopod characters (e.g., many serially similar appendages), and Sida crystallina (Cladocera, Ctenopoda), which likely exhibits plesiomorphic cladoceran traits (e.g., six pairs of serially similar appendages). We combine (semi-)high-speed recordings of behavior with confocal laser scanning microscopy analyses of musculature to infer functionality and homologies of locomotory and filtratory appendages in the two groups. Our morphological study shows that the musculature in all trunk limbs (irrespective of limb size) of both C. hislopi and S. crystallina comprises overall similar muscle groups in largely corresponding arrangements. Some differences between C. hislopi and S. crystallina, such as fewer trunk limbs and antennal segments in the latter, may reflect a progenetic origin of Cladocera. Other differences seem related to the appearance of a specialized type of swimming and feeding in Cladocera, where the anterior locomotory system (antennae) and the posterior feeding system (trunk limbs) have become fully separated functionally from each other. This separation is likely one explanation for the omnipresence of cladocerans, which have conquered both freshwater and marine free water masses and a number of other habitats.     

Feeding Habits of Diaptomus ashlandi and Diaptomus sicilis in Lake Michigan

The herbivorous habits of Diaptomus ashlandi and Diaptomus sicilis were studied at a nearshore station in Lake Michigan. Filtering and feeding rates were measured from December 1975 to October 1976. Feeding was measured from the change in concentration in three size fractions of chlorophyll a. D. ashlandi fed only on particles in the smallest size class, while the larger D. sicilis consumed particles in all three size fractions. During the isothermal period (winter and spring), the feeding rates of both species were proportional to temperature or chlorophyll concentration. D. sicilis preferentially ingested particles in the largest size class, when this size class increased significantly during the spring bloom. During the summer the highest feeding rates of D. ashlandi occurred in the epilimnion. The highest feeding rates of D. sicilis were in the metalimnion, where large particles in a subsurface chlorophyll maximum were available for consumption. These results indicated the importance of the depth distributions of herbivores and phytoplankton for feeding.     

Rosette Recruitment of a Rare Endemic Forb (Gaura neomexicana Subsp. coloradensis) with Canopy Removal of Associated Species

Gaura neomexicana subsp. coloradensis Munz (Onagraceae), (Colorado butterfly plant), a short‐lived perennial forb, became listed as a threatened species by the U.S. Fish and Wildlife Service in October 2000 because of its small global range and population size. This endemic subspecies consists of only 18 extant populations within southeastern Wyoming, southwestern Nebraska, and northeastern Colorado. Wyoming occurrences are on private land with the exception of three occurrences on F. E. Warren Air Force Base, near Cheyenne. Gaura neomexicana subsp. coloradensis may be displaced by many competitors, including Cirsium arvense (L.) Scop. (Canada thistle), an invasive, noxious weed. In June 1998, three G. neomexicana subsp. coloradensis populations were examined for rosette establishment in the presence of and after removal of associated species. The four study treatments removed: (1) Cirsium arvense, (2) grass and forb canopy and associated litter, (3) grass and forb canopy, litter, and C. arvense, and (4) no removal of associated plant species (control). Mature G. neomexicana subsp. coloradensis plants were evaluated for height, number of leaves, and capsule production. The mature plant characteristics monitored in 1998 were not good indicators of subsequent G. neomexicana subsp. coloradensis rosette densities in the following spring. Recruitment was assessed by counting new rosettes in the fall 1999. Although both G. neomexicana subsp. coloradensis and the exotic C. arvense have comparable habitat, removal of C. arvense did not increase G. neomexicana subsp. coloradensis vegetative growth, seed capsule production, or rosette density. Removal of other forbs, grass, and litter, however, increased G. neomexicana subsp. coloradensis rosette density in the following two years, indicating that canopy removal of associated species can enhance rosette establishment of this rare native. The accumulation of dense vegetative cover and litter associated with the absence of herbivory and fire may contribute to the decline of rare species in rich riparian habitats. Return of herbivory and fire in mesic sites to reduce standing biomass accumulations should be considered in restoring recruitment potential to rare monocarpic species.     

Direct evidence of tidally oriented behavior of the copepod <Emphasis Type="Italic">Eurytemora affinis</Emphasis> in the Seine estuary

The calanoid copepod Eurytemora affinis dominates the zooplankton communities in most northern hemisphere estuaries. A recurrent behavior noticed in several estuaries suggests that this species maintains its horizontal position through vertical migration. In order to investigate this behavioral strategy in detail, we sampled E. affinis nauplii, copepodids and adults at high frequency at a fixed point in the low salinity zone of the Seine estuary. We took water samples in sub-surface and near bottom during a 12- and 50-h tide cycle, in May 2002 and May 2004, respectively. Since nauplii have weak swimming capabilities compared to copepodid and adult stages, we proposed here to consider them as passive particles. The variation of the relative abundances of nauplii to older stages regarding the vertical position was therefore viewed as direct evidence of the behavior of later stages. We showed that copepodids and adults tried to stay close to the bottom during the ebb, were re-suspended in the water column 1 h after the low tide during the maximum current velocity, and then vertically migrated back to the bottom layer when current velocity decreased. We also showed that copepodids behaved as adults, with only a lower velocity, taking more time to go near the bottom.     

Impedance traces of copepod appendage movements illustrating sensory feeding behaviour

An experimental system incorporating a computerized micro-impedance unit has been used to make direct measurements of the activity of copepod cephalic appendages. As the appendages are used to both propel the copepod through water and handle particles, it follows that appendage activity reflects feeding behaviour.To investigate the sensory feeding behaviour of copepods, their activity was recorded with food stimuli varying in size and chemical composition. Sample impedance traces are given for the appendage movements of Temora longicornis in the presence of: 1 — filtered seawater; 2 — beads; 3 — phytoplankton cells; 4 — dissolved free amino acids. The normal appendage movements shown in filtered seawater were modified when copepods were offered particles and dissolved chemicals. Results show that chemical and mechanical stimuli are responsible for the recognition and selection of food. Impedance traces distinguish between behavioural responses such as: antennule flicks, leg kicks, combing, handling and rejection of particles. Spectral analyses of traces have demonstrated that differences in beat pattern are significant.Paper presentation, at the Third International Conference on Copepoda. British Museum (Natural History), London, U. K., 10–14 August, 1987.     

Colonization of natural substrata of different roughness and colour by Ephemeroptera nymphs using retrieval and direct observation techniques

We investigated the colonization of stream substrata by Ephemeroptera nymphs in a Canadian Rocky Mountain stream. Stream substrata used in our study were different textures (smooth and rough) and colours (dark and light); the taxonomic groups studied were Drunella coloradensis Dodds (Ephemerellidae), Heptageniidae (Cinygmula and Epeorus) and a Baetis Ameletus group (Baetidae and Siphlonuridae). We tested the hypothesis that as D. coloradensis nymphs and other abundant Ephemeroptera nymphs increase in size they choose a dark substratum to remain cryptic and a substratum of greater texture to reduce the risk of being swept from the substratum by the water flow. Two approaches were used to test this hypothesis, a substratum basket experiment and direct observations.For the substratum basket experiment, there were no statistical differences in abundance of nymphs on the substratum types. But there was a general tendency that the D. coloradensis nymphs were least abundant on the rough dark-coloured substratum and developmental stage three nymphs were most abundant on the smooth white-coloured substratum.Colonization was investigated for other Ephemeroptera nymphs by making direct observations on the substratum baskets using a glass-bottomed box. For the taxonomic groups and two colonization periods, nymphs were statistically at greatest density on the smooth white-coloured substratum and lowest on the smooth black-coloured substratum. The Baetis-Ameletus groups were also found at greatest densities on the rough dark-coloured substratum.     

Nymphal trophic behaviour of two Nemouridae species (Insecta,Plecoptera) in the Curone creek (northern Apennines,Italy)

Feeding habits of the Euholognathan stoneflies Nemoura hesperiae and Amphinemura sulcicollis are studied in a little stream located in northern Italy. The former fed mainly on coarse particulate organic matter (CPOM), together with detritus, while the latter fed principally on detritus and, in a lower concern, on CPOM. Thus they can be catalogued fundamentally as shredder and collector‐gatherer, respectively. Shifts in diet composition were detected in relation to size in both taxa. Probably niche overlapping does not exist despite both species having similar size and nymphal development duration, and coinciding in time. This work supports that, though functional feeding groups are very useful, they should not be inferred from related species or higher taxonomical levels. Even intra‐population differences and ontogenetic shifts should be considered in fine scale studies.     

The Natural Feeding and Resource Partitioning of Erpobdella octoculata L. and Erpobdella testacea SAV. in Lake Esrom,Denmark

The natural feeding and competitive interactions between populations of Erpobdella ostoculata and Erpobdella testacea were investigated in Lake Esrom, Denmark, during 1979/80. Statistical analysis revealed that the size of the leeches was the main factor involved in resource utilization. Size-specific feeding included differences in the numerical composition of diets and the size of individual prey. Five weight classes of Erpobdella spp. were distinguished. The number of prey per leech increased from 1.63-2.84 for the first to the fifth weight class, whereas diversity in feeding averaged 1.71 prey per leech. Quantification of competitive interactions indicated that the temporal differences in weight class composition of the two species were sufficient to explain coexistence, although both species have an annual life cycle, and neither species showed taxonomic food refuges.     

Chromosomal polymorphism in two species of the guarani group of drosophila

Summary Studies of chromosomal polymorphism in natural populations of D. guaramunu and D. griseolineata in Rio Grande do Sul, Brazil, confirmed and extended previous findings in these species. The more common D. guaramunu is much more polymorphic than its closest but less common relative D. griseolineata. In the former species, the frequencies of four inversions showed differences from locality to locality, and five inversions of the same species showed temporal changes in the population of Bexiga. It was not possible to correlate the differences with any particular factor in the environment, though the temporal changes were perhaps due to seasonal climatic cycles.Inversion heterosis was not evident in either species — instead, inversion Ee of D. guaramunu was found to be less frequent than expected on the basis of the Hardy-Weinberg equilibrium in the localities of Emboaba, Bexiga and Feliz. The implications of these findings are discussed.Fellow of the Conselho Nacional de Pesquisas.     

Invader–invader interactions in relation to environmental heterogeneity leads to zonation of two invasive amphipods, Dikerogammarus villosus (Sowinsky) and Gammarus tigrinus Sexton: amphipod pilot species project (AMPIS) report 6

As biological invasions continue, interactions occur not only between invaders and natives, but increasingly new invaders come into contact with previous invaders. Whilst this can lead to species replacements, co-existence may occur, but we lack knowledge of processes driving such patterns. Since environmental heterogeneity can determine species richness and co-existence, the present study examines habitat use and its mediation of the predatory interaction between invasive aquatic amphipods, the Ponto-Caspian Dikerogammarus villosus and the N. American Gammarus tigrinus. In the Dutch Lake IJsselmeer, we found broad segregation of D. villosus and G. tigrinus by habitat type, the former predominating in the boulder zone and the latter in the soft sediment. However, the two species co-exist in the boulder zone, both on the short and longer terms. We used an experimental simulation of habitat heterogeneity and show that both species utilize crevices, different sized holes in a plastic grid, non-randomly. These amphipods appear to optimise the use of holes with respect to their ‘C-shape’ body size. When placed together, D. villosus adults preyed on G. tigrinus adults and juveniles, while G. tigrinus adults preyed on D. villosus juveniles. Juveniles were also predators and both species were cannibalistic. However, the impact on G. tigrinus of the superior intraguild predator, D. villosus, was significantly reduced where experimental grids were present as compared to absent. This mitigation of intraguild predation between the two species in complex habitats may explain the co-existence of these two invasive species.     

Tracking predator density dependence and subterranean predation by carabid larvae on slugs using monoclonal antibodies

1. Subterranean carabid larvae are more numerous than surface‐active adults, yet very little is known about their ecological significance, dietary preferences or ability to regulate populations of prey species, particularly pests. Part of the reason for this is that predator–prey interactions beneath the soil are almost impossible to observe. 2. Extensive field studies have shown that adult Pterostichus melanarius (Illiger) can affect the temporal and spatial dynamics of their slug prey. However, if larvae too are feeding on slugs, this could radically affect overall predator–prey dynamics. 3. We tested the hypotheses that P. melanarius larvae would kill and consume two slug species, Deroceras reticulatum Müller and Arion intermedius Normand, under laboratory and semi‐field conditions, and that there would be no significant difference in rates of predation on these slug species. 4. A new monoclonal antibody was developed that was capable of detecting the presence of slug proteins in the guts of P. melanarius larvae. 5. Pterostichus melanarius larvae killed both A. intermedius and D. reticulatum in the laboratory, feeding to a greater extent, and growing more rapidly, on the latter. The larvae were equally effective at reducing numbers of both slug species in a crop of wheat grown in semi‐field mini plots, but predation was affected by density‐dependent intra‐specific competition amongst the beetle larvae. 6. Future modelling of the dynamic interactions between carabids and slugs will need to take into account predation by larvae.     

中国毛茛科翠雀属的分类学研究(十三):狭序翠雀花的名实订正

发现根据甘肃南部标本描述的毛茛科狭序翠雀花（Delphinium angustipaniculatum W.T.Wang）与此前记载分布于甘肃南部和四川北部的松潘翠雀花（D.sutchuenense Franch.）属于同一种植物,故将前者处理为后者的异名。讨论了松潘翠雀花与其可能的近缘种秦岭翠雀花（D.giraldii Diels）和疏花翠雀花（D.sparsiflorum Maxim.）的形态区别。     

Some Consequences of Size Variability in Juvenile Prickly Sculpin, Cottus asper

Variation in size among fishes can have important ecological consequences. The origin and persistence of variation in size among a cohort of juvenile prickly sculpin, Cottus asper, was explored experimentally by manipulating the size variability in juvenile fish and the presence of conspecific adult fishes in a factorial design. I found that adult fish significantly suppressed the growth and survivorship of juveniles, while the variance treatment alone had no effect on growth or survivorship. The presence of adults was also associated with a decrease in size variability in the high variance treatment. An analysis of juvenile growth patterns revealed no evidence for size dependent growth or temporal correlations in growth (`growth autocorrelation'), and are consistent with a lack of amplification of size variance through time. Juvenile C. asper appear to have been feeding on a relatively homogeneous resource and utilizing those resources similarly. Thus, patterns of individual growth and the presence of adult conspecifics may both serve to dampen size differences in this coastal fish species.     

Limits to genetic bottlenecks and founder events imposed by the Allee effect

The Allee effect can result in a negative population growth rate at low population density. Consequently, populations below a minimum (critical) density are unlikely to persist. A lower limit on population size should constrain the loss of genetic variability due to genetic drift during population bottlenecks or founder events. We explored this phenomenon by modeling changes in genetic variability and differentiation during simulated bottlenecks of the alpine copepod, Hesperodiaptomus shoshone. Lake surveys, whole-lake re-introduction experiments and model calculations all indicate that H. shoshone should be unlikely to establish or persist at densities less than 0.5–5 individuals m⁻³. We estimated the corresponding range in minimum effective population size using the distribution of habitat (lake) sizes in nature and used these values to model the expected heterozygosity, allelic richness and genetic differentiation resulting from population bottlenecks. We found that during realistic bottlenecks or founder events, >90% of H. shoshone populations in the Sierra Nevada may be resistant to significant changes in heterozygosity or genetic distance, and 70–75% of populations may lose <10% of allelic richness. We suggest that ecological constraints on minimum population size be considered when using genetic markers to estimate historical population dynamics.     

The microcopepod fauna in the Eastern Mediterranean and Arabian Seas: a comparison with the Red Sea fauna

Multiple opening-closing nets of 0.05 mm mesh size were employed to study the community structure and vertical distribution of microcopepods at selected stations in the Red Sea, Arabian Sea and Eastern Mediterranean Sea down to a maximum depth of 1850 m. Calanoids, cyclopoids (Oithona and Paroithona) and poecilostomatoids (mainly Oncaea) were the 3 most abundant orders. In the epipelagic zone (0–100 m), these orders occurred at similar abundance levels, whereas in the meso- and bathypelagic zones the poecilostomatoid genus Oncaea dominated numerically by about 60–80% of all copepodids.The species diversity of Oncaea in the Red Sea is compared with preliminary results from the two adjacent regions. In the deep Eastern Mediterranean Sea, the number of species appears to be similar to that in the deep Red Sea and low as compared to the deep Arabian Sea. In this latter area an extremely speciose Oncaea fauna was found at depth below the oxygen-minimum-zone (900–1850 m). The results are related to the differences in the hydrographic conditions of these 3 areas.     

Ontogeny of pistillate flowers and inflorescences in Nothofagus subgenus Lophozonia (Nothofagaceae)

  Cupule and pistillate inflorescence morphology is described in detail for the first time for a number of species in Nothofagus subgenus Lophozonia. The architecture and temporal appearance of prophylls, flowers, and cupule valves is dichasial; the cupule valves are third and higher order shoot systems. Comparative studies demonstrate that the dorsal rows of appendages on the cupule valves have a similar arrangement, and each row exhibits the same temporal pattern of development. This suggests that the cupule valves in all species have a modular construction and consist of rows of homologous appendages. This study clearly demonstrates the uniform groundplan of the pistillate inflorescence in this subgenus. Received March 14, 2002; accepted May 8, 2002 Published online: September 13, 2002     

EVOLUTIONARY GENETICS OF TWO SIBLING SPECIES,DROSOPHILA SIMULANS AND D. SECHELLIA

Drosophila simulans and D. sechellia are sibling species, the former cosmopolitan and the latter restricted to the Seychelles Islands. We used classical genetic analysis to measure the numbers and effects of genes responsible for reproductive isolation and morphological differences in male genitalia between these species. At least five loci are responsible for male sterility in hybrids, with the strongest effects produced by at least two genes on the X chromosome. At least three (and probably four) loci are responsible for the interspecific difference in the size of the posterior process of the male genital arch. These genetic results, as well as the pattern of morphological divergence between the species, show several parallels with the divergence between D. simulans and its other island relative, D. mauritiana. We also present the DNA sequence of a 4.5 kilobase region containing the alcohol dehydrogenase (Adh) locus of D. sechellia, and combine this with previous data to reconstruct the phylogenies of the three species and their more distant relative D. melanogaster. Both D. mauritiana and D. sechellia are very closely related to D. simulans. Although most phylogenies show the two island species to be independent offshoots of the D. simulans lineage (with D. sechellia the more recent), the branch points are too close to make this conclusion unambiguous. The genetic and evolutionary parallels between the simulans/mauritiana and the simulans/sechellia divergences may therefore represent either a striking evolutionary convergence or a close common ancestry of the island species. A comparison of Adh alleles within species shows that the divergence among them may be almost as large as among alleles from different species. We conclude that many of the nucleotide differences among these species actually represent polymorphisms within common ancestors. It may be difficult to build accurate phylogenies using only a single DNA sequence from each species.     

Opportunistic or selective? Stage-dependent feeding behavior in a wild vector of Chagas disease

The composition and contribution of different host species in the dynamics of vector-borne zoonotic parasites are particularly relevant for public health. Hence, the study of host selection by vectors is fundamental. Developmental stage and infection status are factors that may modulate vector feeding behavior. In the semi-arid Mediterranean ecosystem of South America, the transmission of Trypanosoma cruzi, the protozoan causing Chagas disease, includes the triatomine vector Mepraia spinolai and several vertebrate species. In this field study, we examined whether M. spinolai exhibits an opportunistic feeding behavior dependent upon developmental stage and/or infection status. We found that M. spinolai does not feed according to the relative availability of vertebrate species. In addition, early stage nymphs (first/second instars) fed on twice as many different species as middle (third/fourth instars) and late (fifth instars and adults) M. spinolai, with the former feeding on native rodents and lizards and the latter mostly on rabbits. Infected and uninfected M. spinolai showed similar feeding profiles. Wild triatomine species might be described as stage-dependent selective blood feeders, as a consequence of the temporal and spatial scale at which host-vector interactions occur, highlighting that all developmental stages might be infected and capable of transmitting T. cruzi.     

Effects of seasonal variation in host quality and availability on parasitism by the egg parasitoid Telenomus coloradensis

Hemlock looper, Lambdina fiscellaria (Guenée) (Lepidoptera: Geometridae), is one of the most important defoliator in North American forests. The common egg parasitoid Telenomus coloradensis Crawford (Hymenoptera: Platygastridae) plays a significant role as a natural control agent, with parasitism levels in spring typically higher than in fall. The objectives of this study were to quantify changes in host acceptance and reproductive performance of the parasitoid in relation to (1) host egg fertilization in fall, (2) host diapause status, (3) host embryonic development in spring, and (4) host deprivation during summer. Our results indicate that T. coloradensis do not have the capacity to develop in unfertilized host eggs, whereas early‐diapausing eggs are more suitable for the parasitoid than post‐diapausing eggs. Furthermore, the host physiological suitability decreases with embryonic development in spring. Finally, a host deprivation period during the summer tends to negatively affect the parasitic activity of T. coloradensis. These laboratory results confirm previous hypotheses concerning T. coloradensis seasonal ecology and contribute to a better understanding of the effect of hemlock looper egg physiology and availability on the reproductive potential of T. coloradensis.