Die Antennen-Putzeinrichtung der Adephaga (Coleoptera), parallele evolutive Vervollkommnung einer komplexen Struktur

The antenna cleaner of the Adephaga (Coleoptera), parallel evolutionary perfection of a complex structure Independent evolutionary perfection of a complex character ist to be expected in a splitting phylogenetic line, if selective advantages for improving this character persist in the various lines after splitting. Independent perfection leading to very similar results is termed “parallel perfection”. It can either be based on a narrow channelling, due to limitations of the evolutionary substrate (i. e. morphological structures and patterns of behaviour, inherited from the common ancestor), or on very specific functional advantages. The evolution of the antenna cleaner on the protibia of adephagous beetles is analysed under this aspect. The phylogenetic relationships within the Geadephaga are discussed. After defining the terms “anisochaetous” and “isochaetous” somewhat differently from previous usage, the two phylogenetic lines Anisochaeta (Carabidae in a broad sense) and Isochaeta (Metriidae, Ozae-nidae, Paussidae) may be considered sister groups. The Cicindelidae might have split off from the line of the Anisochaeta very early. The Rhysodidae are interpreted as an aberrant offshoot from the primitive adephagous stock. An attempt is made to reconstruct the antenna cleaner of the common ancestor of Anisochaeta and Isochaeta from the sum of plesiomorphous characters. This primitive cleaner might have corresponded in many respects to the array of setae found on an undifferentiated protibia (see comparison with mesotibia). The cleaner of the Cicindelidae, in spite of many plesiomorphous characters cannot be regarded as a model for the common ancestral form, because it already shows some differentiation towards the condition found in the Anisochaeta. The antenna cleaners of highly evolved Anisochaeta and Isochaeta, which correspond surprisingly well in many details, can be traced back to a common basis, but most of their conformities must have evolved independently. The following parallelisms were established: the ventral side of the tibia is shifted to the functional inner side and forms an oblique groove (cleaning channel); the antenna cleaning comb, originally placed transversely, becomes lengthwise oriented and is shifted in proximal direction; several setae are modified so as to form together a pressure clamp; the cleaning channel is completed by a spine-like extension projecting from the tibia; the terminal setal ring forms an additional comb. The gradual perfection of the antenna cleaner can be studied in the more primitive species in both the groups Anisochaeta and Isochaeta. Apparently, the two lines entered different pathways of perfection to begin with, and the cleaners became similar again by convergence later on. This indicates a very slight channelling effect of the common evolutionary substrate. The numerous conformities in the higher evolved genera must result from “parallel selection” (channelling for functional reasons). By observing grooming behaviour in primitive and highly evolved genera, the following advantages connected with the more complex structures could be established: the engagement of the antenna into the cleaning channel is facilitated; the mechanical strain on the antenna during grooming is reduced; the cleaning of antennae and eyes becomes more effective. That the antenna cleaner itself can better be kept clean by a comb of the Mesotibia was proved quantitatively.     

Functional morphology of amplexus (clasping) in spinicaudatan clam shrimps (Crustacea,Branchiopoda) and its evolution in bivalved branchiopods: A video‐based analysis

Male clam shrimps (Crustacea: Branchiopoda: Laevicaudata, Spinicaudata, and Cyclestherida) have their first one or two trunk limb pairs modified as “claspers,” which are used to hold the female during mating and mate guarding. Clasper morphology has traditionally been important for clam shrimp taxonomy and classification, but little is known about how the males actually use the claspers during amplexus (clasping). Homologies of the various clasper parts (“movable finger,” “large palp,” “palm,” “gripping area,” and “small palp”) have long been discussed between the three clam shrimp taxa, and studies have shown that only some structures are homologous while others are convergent (“partial homology”). We studied the clasper functionality in four spinicaudatan species using video recordings and scanning electron microscopy, and compared our results with other clam shrimp groups. General mating behavior and carapace morphology was also studied. Generally, spinicaudatan and laevicaudatan claspers function similarly despite some parts being nonhomologous. We mapped clasper morphology and functionality aspects on a branchiopod phylogeny. We suggest that the claspers of the three groups were adapted from an original, simpler clasper, each for a “stronger” grip on the female's carapace margin: 1) Spinicaudata have two clasper pairs bearing an elongated apical club/gripping area with one setal type; 2); Cyclestherida have one clasper pair with clusters of molariform setae on the gripping area and at the movable finger apex; and 3) Laevicaudata have one clasper pair, but have incorporated an additional limb portion into the clasper palm and bear a diverse set of setae. J. Morphol. 278:523–546, 2017. © 2017 Wiley Periodicals, Inc.     

Seasonal Development of Phytoplankton in Fish Ponds

At the fish ponds under study the authors defined several types of plankton which have been frequently found during the season (April – October). These types are: Early-spring maximum of phytoplankton, Depression of phytoplankton, Bloom of Aphanizomenon, Maximum of Chlorococcales. The periods of “depression” seem to be typical for the managed carp ponds in the spring. They are characterized by the low density of rapidly reproducing algal populations (e. g. Cryptomonas) and by the dense populations of large cladocerans of the genus Daphnia. Chlorophyll in phytoplankton is less than 5μg/l, transparency is higher than 2 meters. Periods of the spring depression may be followed by the maxima of either Aphanizomenon or Chlorococcales, with concentrations of chlorophyll increasing to 100 μg/l and more. The change from the phase of “depression” to the “maximum of Chlorococcales” is accompanied by decrease in numbers of Daphnia and increase in numbers of the small cladoceran species, but all the mechanisms responsible for the transition are not yet fully understood.     

Primary homology assessment of structures in the female atrial system among species of the Polycystididae (Rhabditophora, Eukalyptorhynchia)

Abstract. The female atrial system of members of the Polycystididae has been studied at the light microscopical level and compared among the constituent taxa. Based on the criteria of position and conjunction, hypotheses of homology are put forward and compared with the assessments of homology found in earlier literature. Contradictory terminology is synonymized and adapted to fit new findings that recognize homology among structures in different taxa. Based on differences in position and structure, 2 types of female duct are recognized: female duct type I and female duct type II. The term “female bursa” is restricted to a sperm resorbing organ at the proximal end of the female duct type I and/or at the end of a common oviduct. Some species have bundles of glands at the place where the oviduct(s) enter the female duct type I, which are considered homologous among these species. Different types of seminal receptacles are recognized. The term “insemination duct” is restricted to the ducts which in some species, in addition to the oviducts, connect the ovaries with the female duct type I. A single muscular duct, the common oviduct, connects the joined oviducts with the female duct type I in species of Duplacrorhynchus and is considered homologous with the similar duct present in some other species. A uterus is always present, entering the common genital atrium through its frontal wall, or entering a female duct type II. This morphological review of the female atrial system reveals a mosaic distribution of characters if applied to previous phylogenies that are based primarily on the male reproductive system.     

Anthessius kimjensis,a new species of Anthessiidae (Copepoda; Poecilostomatoida) associated with the pelecypod Solen grandis Dunker in Korea

Anthessius kimjensis n. sp. is described based on the specimens recovered from the pelecypod Solen grandis Dunker in the Korean coast of the Yellow Sea. The new species is distinguished from congeners by having a combination of characters: setal formula II, I, 5 on distal segment of leg 4 exopod, three terminal hooks on distal segment of second antenna, and ratio of length to width on caudal ramus (2.2:1).     

Spirometra species from Asia: Genetic diversity and taxonomic challenges

Despite considerable controversy concerning the taxonomy of species within the genus Spirometra, human sparganosis and spirometrosis mainly in Asia and Europe has long been confidently ascribed to Spirometra erinaceieuropaei. Recently, the mitochondrial genomes of purported “S. erinaceieuropaei”, “Spirometra decipiens” and “Spirometra ranarum” from Asia have been determined. However, it has been pointed out that the morphological criteria used for identifying these species are unsuitable and thus these identifications are questionable. In the present study, therefore, Spirometra samples from Asia were re-examined based on mitochondrial cytochrome c oxidase subunit 1 gene sequences and the identification of these species was discussed. Haplotype network and phylogenetic analyses revealed that: i) two distinct Spirometra species, Type I and Type II, are present in Asia and neither of which is close to likely European “S. erinaceieuropaei”; ii) Type I is genetically diverse and widely distributed, however Type II is known so far from Japan and Korea; iii) “S. decipiens” and “S. ranarum” reported from Asia are conspecific with Type I; iv) Type I is probably conspecific with Spirometra mansoni, and Type II may represent an undescribed species.     

Detecting shifts in diversification rates without fossils

Studies of shifts in diversification rates and adaptive radiations are difficult when there are no fossils because past events cannot be inferred. The phylogenies of recent species, however, allow one to infer the patterns of past diversifications. I present a new method for estimating the diversification rate of a lineage, provided that a phylogeny of recent species, constructed, for instance, with molecular data, is available. This method was inspired by survival models and takes into account species that are not included in detailed phylogenetic data, provided that approximate dates of origin of these species are known. Likelihood ratio tests and Akaike Information Criterion make it possible to test for differences in diversification among lineages or groups of lineages and, thus, to evaluate adaptive radiation hypotheses. The present modeling approach can easily be extended to include temporal variations in diversification rates. A simulation study showed that the method is statistically consistent, avoiding Type I and Type II errors, and that it is robust to periodic or random fluctuations in the speciation rate. An example is presented with a composite phylogeny of primates.     

Characterization of topological keystone species: Local,global and “meso-scale” centralities in food webs

An important question in the network representation of ecological systems is to determine how direct and indirect interactions between species determine the positional importance of species in the ecosystem. Here we present a quantitative analysis of the similarities and differences of six different topological centrality measures as indicators of keystone species in 17 food webs. These indicators account for local, global and “meso-scale” – intermediate between local and global – topological information about species in the food webs. Using factor analysis we shown that most of these centrality indices share a great deal of topological information, which range from 75% to 96%. A generalized keystone indicator is then proposed by considering the factor loadings of the six-centrality measures, which contains most of the information encoded by these indices. However, the individual ordering of species according to these criteria display significant differences in most food webs. We simulate the effects of species extinction by removing species ranked according to a local and a “meso-scale” centrality indicator. The differences observed on three network characteristics – size, average distance and clustering coefficient of the largest component – after the removal of the most central nodes indicate that the consideration of these indices have different impacts for the ranking of species with conservational biology purposes. The “meso-scale” indicator appears to play an important role in determining the relative importance of species in epidemic spread and parasitism rates.     

The reality of the higher taxonomic categories

A higher category is the result of a population's breaking-throuh into a new adaptive zone (attaining a new grade), and the subsequent adaptive radiation tiere. The taxonomic level of the new group is determined by the amount (quantitative and ualitative) of radiation that occurs; and tgis amount in turn is determined by the number of open sub-zones in the new zone and by the genetic plasticity of the new population. The more the new adaptive zone differs from the old one, the less likely a new population can survive in it. Should such survival occur, however, adaptations of the new population to the new zone will appear quickly. The new population will rapidly become quite different from the old one, minimizing the likelihood of finding intermediate forms, i. e., “missing links.” In addition, the intermediate population – the early entrants into the new zone – will be less well adapted to it than their descendents, and will become extinct. The combination of rapid extinction and an ephemeral transitional population explains the apparent gaps between higher-category taxa. These ideas do not help in solving such dilemmas of taxonomists as equivalence of taxa in different groups, and the taxonomic level at which to place a given group of species. But a higher taxonomic category itself is the result of a very real biological phenomenon, and that fact must influence systematicists.     

Antenna replacement in the evolutionary origin of chloroplasts

The endosymbiotic origin of chloroplasts from unicellular cyanobacteria is presently beyond doubt. Oxygenic photosynthesis is based on coordinated action of the two photosystems (PS), PS I and PS II, cooperating with several variants of the pigment antenna. In cyanobacteria, red algae, and glaucophytes, phycobilisomes (PBS) act as antennae, while in terrestrial plants, as well as in most macro- and microalgae, antennae are formed by chlorophyll a/b- and chlorophyll a/c-containing proteins. Advantages and disadvantages of the PBS antenna compared to other light-harvesting complexes form the basis for adaptive variations of the antenna in the course of development of eukaryotic photosynthesis. During the evolution of the “green” and “chromophyte” lineages of the chloroplasts, PBS, in spite of their optimal features of light absorption, were replaced by chlorophyll a/b- and chlorophyll a/c-containing light-harvesting complexes. Development of the cell wall associated with the limitation of motility and tissue formation in photosynthetic eukaryotes were the factors responsible for the antenna shift. The subsequent redistribution of cell resources in favor of cellulose biosynthesis required for increased CO₂ consumption, higher PS II levels, and greater number and density of the thylakoids in the chloroplasts, was incompatible with the energy-consuming and overly large PBS antenna.     

The ontogenetical and supposed phylogenetical fate of the parietal muscles in the Ctenostomata (Bryozoa)

A new term, “trophon”, has been introduced for the feeding and sexually propagating unit in the stolonate ctenostomes; the “stolon” and the “trophon” represent “sukzooids”, because they are homologous only to parts of the autozooids in the uniserially arranged ctenostomes. – The ontogeny of the muscles, especially the parietal muscles, has been studied in 17 species of ctenostomatous bryozoans belonging to different sub-groups of this order; the main effort was directed toward the details of the differentiations in living animals (14 of the investigated species). In various species of the Ctenostomata two separate sets of parietal muscles develop – probably as a consequence of the elongation of the cystid; the functional parietal muscles in the trophons of the stolonial forms are homologous to the secondary parietal muscles found in several uniserial ctenostomes. Contrary to the supposition by Soule (1954) in the different families of the serially arranged forms the muscles appear in different sequences during the development of the zooid and partly have different ontogenetical fates. Neither the “Carnosa” nor the “Paludicellea” (s. 1.) nor the “Stolonifera” can be maintained as taxa with the argument of a similar ontogeny of the muscles. A detailed comparison regarding the fate of the parietal muscles (and the polypide anlage) showed similarities between certain of the uniserial to either some of the stolonial, the sheet like, or the cystid-fusing forms suggesting a separate evolutionary origin of the advanced forms in those groups of the serially growing forms. – A new family, Pottsiellidae fam. nov., has been introduced and defined.     

Two roles for ecological surrogacy: Indicator surrogates and management surrogates

Ecological surrogacy – here defined as using a process or element (e.g., species, ecosystem, or abiotic factor) to represent another aspect of an ecological system – is a widely used concept, but many applications of the surrogate concept have been controversial. We argue that some of this controversy reflects differences among users with different goals, a distinction that can be crystalized by recognizing two basic types of surrogate. First, many ecologists and natural resource managers measure “indicator surrogates” to provide information about ecological systems. Second, and often overlooked, are “management surrogates” (e.g., umbrella species) that are primarily used to facilitate achieving management goals, especially broad goals such as “maintain biodiversity” or “increase ecosystem resilience.” We propose that distinguishing these two overarching roles for surrogacy may facilitate better communication about project goals. This is critical when evaluating the usefulness of different surrogates, especially where a potential surrogate might be useful in one role but not another. Our classification for ecological surrogacy applies to species, ecosystems, ecological processes, abiotic factors, and genetics, and thus can provide coherence across a broad range of uses.     

Interspecific,ontogenetic, and sexual variation in ozopore morphology among cosmetid harvestmen (Arachnida,Opiliones, Laniatores)

The ozopores of cosmetid harvestmen rest upon lateral projections of the carapace, have simple or highly reduced channels, and are partially obscured by enlarged dorsal processes associated with coxae I and II. Rather than use scent gland secretions to form a chemical shield on the dorsum, the cosmetid harvestman exhibits a unique defensive behavior known as “leg dabbing” in which the distal tip of tarsus I or II is dipped into fluid that accumulate at the base of coxa II and the droplet on the tarsus is pointed toward the predator. Relatively little is known about interspecific variation in ozopore morphology among cosmetid harvestmen. In this study, we used scanning electron microscopy to examine the ozopores of males and females of nine species as well as those of antepenultimate nymphs for two species. Among adults, we found differences between species in the shapes of the ozopores (round or subtriangular), the morphology of the dorsal and lateral channels (if present), and the relative size, shape and armature of the dorsal posterior process (dpp) of coxa I and the dorsal anterior process (dap) of coxa II. Our observations suggest that the morphology of dpp I and dap II could be sources for systematic characters in future phylogenetic studies of the Cosmetidae. We observed ontogenetic differences but relatively little intersexual variation in the morphology of the ozopore. The ozopores of nymphs are generally more oval than those of adults and the opening of the ozopore of the nymph is less obstructed, if at all, by the dorsal coxal processes of legs I–II. These morphological differences suggest that nymphs may use scent gland secretions in a manner different from that of adults.     

ECOLOGICAL OPPORTUNITY AND THE RATE OF MORPHOLOGICAL EVOLUTION IN THE DIVERSIFICATION OF GREATER ANTILLEAN ANOLES

The pace of phenotypic diversification during adaptive radiation should decrease as ecological opportunity declines. We test this prediction using phylogenetic comparative analyses of a wide range of morphological traits in Greater Antillean Anolis lizards. We find that the rate of diversification along two important axes of Anolis radiation—body size and limb dimensions—decreased as opportunity declined, with opportunity quantified either as time elapsed in the radiation or as the diversity of competing anole lineages inferred to have been present on an island at different times in the past. Most previous studies of the ecological opportunity hypothesis have focused on the rate of species diversification; our results provide a complementary perspective, indicating that the rate of phenotypic diversification declines with decreasing opportunity in an adaptive radiation.     

Flagellar setae of the second antennae in decapod shrimps: sexual dimorphism and possible role in detection of contact sex pheromones

Summary

The antenna 2 (antennal) flagella of decapod shrimps are chemotactile, and their setae are proposed as sensilla involved in recognition of females by males via a contact sex pheromone on the surface of the female. Male recognition of females receptive to mating occurs in many caridean species upon contact of male antennal flagella with the surface of a newly molted parturial female. The hypothesis of sexual dimorphism in the number and kind of setae on the antennal flagella of four caridean and one penaeid shrimp species was tested with setal counts and observations on setal morphology. Unique male antennal setae (“male-specific sensilla”) were not identified in any of the species investigated. However, the abundance of antennal setae was significantly greater in males than in breeding females in the palaemonid carideans Palemonetes pugio and Macrobrachium ohione. In the hippolytid caridean Thor manningi and alpheid caridean Alpheus normanni, no sexual dimorphism in setal abundance was demonstrated. In the penaeoid Rimapenaeus similis, males had a higher abundance of antennal setae than the larger breeding females but so did juvenile females, similar in size to males. The sexual dimorphism in antennal sensilla in the palaemonid species and its absence in A. normanni might be related to their different mating systems, but no such association is suggested for T. manningi and R. similis. Setal morphology suggestive of chemoreceptive function (a terminal pore) was observed in all species.     

The cryptic origins of evolutionary novelty: 1000‐fold faster trophic diversification rates without increased ecological opportunity or hybrid swarm

Ecological opportunity is frequently proposed as the sole ingredient for adaptive radiation into novel niches. An additional trigger may be genome‐wide hybridization resulting from “hybrid swarm.” However, these hypotheses have been difficult to test due to the rarity of comparable control environments lacking adaptive radiations. Here I exploit such a pattern in microendemic radiations of Caribbean pupfishes. I show that a sympatric three species radiation on San Salvador Island, Bahamas diversified 1445 times faster than neighboring islands in jaw length due to the evolution of a novel scale‐eating adaptive zone from a generalist ancestral niche. I then sampled 22 generalist populations on seven neighboring islands and measured morphological diversity, stomach content diversity, dietary isotopic diversity, genetic diversity, lake/island areas, macroalgae richness, and Caribbean‐wide patterns of gene flow. None of these standard metrics of ecological opportunity or gene flow were associated with adaptive radiation, except for slight increases in macroalgae richness. Thus, exceptional trophic diversification is highly localized despite myriad generalist populations in comparable environmental and genetic backgrounds. This study provides a strong counterexample to the ecological and hybrid swarm theories of adaptive radiation and suggests that diversification of novel specialists on a sparse fitness landscape is constrained by more than ecological opportunity and gene flow.     

What can resting egg banks tell about cladoceran diversity in a shallow subtropical lake?

Dormant stages (“resting eggs”) produced by cladocerans can persist for long periods of time in sediments and restore populations once the environmental conditions become favorable again. Lake Blanca, a subtropical shallow eutrophic lake, hosts a cladoceran community dominated by small-sized species. Previous studies on zooplankton resting eggs suggested that the cladoceran genera Daphnia and Simocephalus were present, but they had never been found before in water samples. In the present study, we compared a biweekly active community sampling with the resting egg bank (passive cladoceran community) from littoral and pelagic zones. Moreover, we tested the amount of samples required to have a representative reconstruction of the diversity in both compartments (water and sediment). Lake Blanca showed a relatively high cladoceran species richness (24) in the water column, with rapid temporal replacement. Several species were present in water samples during short temporal windows; therefore, to detect these species a strong sampling effort in terms of temporal frequency and spatial distribution was required. Contrary to our expectations, resting egg community showed a lower diversity than the active community; however, we demonstrated that the analysis of resting egg bank composition can help detect general community structure patterns.     

Major radiation of cheilostome bryozoans: Triggered by the evolution of a new larval type?

A macroevolutionary model is developed to account for the “adaptive radiation”; of cheilostome bryozoans that commenced in the Cenomanian after a long phase of low diversity. Living cheilostome species possess one of two types of larvae; planktotrophic (cyphonautes) larvae of relatively long duration, and brooded non‐planktotrophic (coronate) larvae of short duration. Planktotrophic larvae characterize the paraphyletic “malacostegans”; from which “advanced”; cheilostomes with non‐planktotrophic larvae are thought to have evolved monophyletically. Research on other marine invertebrates suggests that gene flow within and between populations is likely to be poorer in species having non‐planktotrophic larvae, and hence the frequency of allopatric and quasi‐sympatric speciation may be greater. Skeletal evidence of larval brooding in the cheilostomes first appears in the late Albian, immediately before their adaptive radiation, and the evolution of non‐planktotrophy with associated increase in speciation rate is proposed to have triggered this radiation.     

Biotech News

Awards: Smoluchowski Award for Suwan Jayasinghe – EMBO recognizes talented young group leaders in Europe New on the market: Full automation from library samples to assay plates with “Calli-L” – New bottles for laboratory use From our sister journals: – Estrogen activity biosensor – Two-photon polymerization for tissue scaffolds – Algae, do not shear this time! – Plant based Plasmodium vaccine – The Global HIV Vaccine Research Cryorepository – Bioreactor for tissue engineered arteries – Nano-magnetic bacteria separation – Predictive biodegradation models – Most read in BTJ: Gene targeting from laboratory to livestock BioEssays highlights: What doesn't kill me... – Time for adaptive radiation to move on? – Looking again (carefully) at thalidomide     

Characterizing the multivariate physiogenomic response to environmental change

Global change is altering the climate that species have historically adapted to – in some cases at a pace not recently experienced in their evolutionary history – with cascading effects on all taxa. A central aim in global change biology is to understand how specific populations may be “primed” for global change, either through acclimation or adaptive standing genetic variation. It is therefore an important goal to link physiological measurements to the degree of stress a population experiences (Annual Review of Marine Science, 2012, 4, 39). Although “omic” approaches such as gene expression are often used as a proxy for the amount of stress experienced, we still have a poor understanding of how gene expression affects ecologically and physiologically relevant traits in non‐model organisms. In a From the Cover paper in this issue of Molecular Ecology, Griffiths, Pan and Kelley (Molecular Ecology, 2019, 28) link gene expression to physiological traits in a temperate marine coral. They discover population‐specific responses to ocean acidification for two populations that originated from locations with different histories of exposure to acidification. By integrating physiological and gene expression data, they were able to elucidate the mechanisms that explain these population‐specific responses. Their results give insight into the physiogenomic feedbacks that may prime organisms or make them unfit for ocean global change.