A eucrustacean from the Cambrian ‘Orsten’ of Sweden with epipods and a maxillary excretory opening

The Cambrian species Paulinecaris siveterae n. gen. n. sp., known from two trunk fragments, represents the first record of epipods (serving as gills and osmoregulatory structures) in a crustacean from the Swedish ‘Orsten’. Moreover, it is the first report of the maxillary excretory opening of a crustacean based on Cambrian material of ‘Orsten’‐type preservation. One specimen comprises the maxillary segment with an appendage and several thoracic segments with parts of their limbs; a second specimen is a fragment possibly of a more posterior part of the trunk. As in other known small eucrustaceans, the tergites of the new species lack prominent tergopleurae, so that the limbs insert directly ventral to the tergal margins. Limb preservation includes the maxilla and several thoracopods, all possessing a prominent, fleshy basipod with six setose endites along their median rim distally to the proximal endite. The presence of long and prominent limbs of P. siveterae suggests that it had good swimming ability, while the slight C‐like curvature of their basal limb part, basipod, indicates involvement of the limbs also in so‐called ‘sucking chambers’ for suspension feeding coupled with locomotion. The estimated total length of P. siveterae, 2–3 mm, is comparable to that of extant cephalocarids, but its appendages are twice as long and wide. The limbs of P. siveterae also differ in size and armature from extant eucrustaceans as well as early representatives of this group known from the ‘Orsten’ assemblages. The general morphology of the limbs, for example in having a fleshy and C‐shaped basipod with several setae‐bearing endites medially, identifies P. siveterae as an entomostracan eucrustacean, but a lack of further details precludes its affinity with any of the in‐group taxa. Three epipods on the outer edge of the basipod, as in P. siveterae, are also known from the Cambrian eucrustacean Yicaris dianensis from China and early ontogenetic stages of extant fairy shrimps (Anostraca); their adult stages have two epipods. This hints at an original number of three epipods in the ground pattern of Entomostraca, but some uncertainty remains with regard to the eucrustacean ground pattern because Malacostraca possess a maximum number of two.     

Tardigrades as ‘Stem-Group Arthropods’: The Evidence from the Cambrian Fauna

The Cambrian fauna can now reasonably be seen as containing many taxa that lie in the stem-groups of the extant phyla. As such, these fossils suggest how both the ‘body plans’ of extant phyla were assembled, and also how various ‘minor’ phyla relate to the larger groupings of today such as the arthropods and annelids.

The various arthropod and lobopod taxa of the Cambrian faunas have been controversial and have generally been considered either as lying in the crown or (occasionally) stem groups of the euarthropods, onychophorans and tardigrades. However, phylogenetic analysis strongly suggests that many of even the most euarthropod-like taxa do not lie within the euarthropod crown-group but are more basal. Further, the commonly expressed view that Cambrian lobopods are in effect stem- or crown-group onychophorans also seems not to be well supported. Lobopods in the Cambrian appear to be diverse and not particularly closely related to one another, and certainly cannot be combined in a monophyletic clade.

Both these advances offer hope that the tardigrades (placed as the sister group to the euarthropods in many analyses of extant taxa, here collectively named the Tactopoda) may be more closely related to some of these Cambrian taxa than others. The challenge for both neontologists and palaeontologists is to refine the systematic analysis of both living and fossil taxa in order to maximise the usefulness of the (admittedly few) characters that unite tardigrades to their Cambrian forbears.     

Exceptionally well‐preserved isolated eyes from Cambrian ‘Orsten’ fossil assemblages of Sweden

Abstract: Eyes other than those of trilobites are rarely preserved in the fossil record. We describe here a set of six tiny, isolated, three‐dimensionally preserved compound eyes. These secondarily phosphatized eyes were etched from ‘Orsten’ limestone nodules dated to the Agnostus pisiformis Biozone from the Cambrian Alum Shale Formation of Sweden. The ovoid eyes arise from an elongated stalk, their surface being covered by a mosaic of regular and hexagonal‐shaped facets representing the surface of ommatidia. Facet size and pattern change within the same specimen from the posterior to the anterior end. With regard to some morphological criteria, we grouped the material in two different morphotypes, type A and B, the first being represented by specimens of two different developmental stages. From stage to stage, mostly growth in overall size and addition of new ommatidia was noticed. Among the meiobenthic ‘Orsten’ arthropods, only the crustacean Henningsmoenicaris scutula has been described as possessing stalked eyes, but the eyes of the largest specimen with preserved eyes of this species are much smaller than the new eyes and do not display any kind of ommatidia on their visual surface. However, fragments of larger specimens of H. scutula and the co‐occurrence of this species with the new isolated eyes in the sieving residues make it likely that the latter belong to this species but belong to more advanced stages than those described previously of H. scutula. Ontogenetically, the eye stalks of this fossil crustacean elongate progressively, while the regular hexagonal facets, lacking in early stages, appear later on.     

The AAT1 locus is critical for the biosynthesis of esters contributing to ‘ripe apple’ flavour in ‘Royal Gala’ and ‘Granny Smith’ apples

The ‘fruity’ attributes of ripe apples (Malus × domestica) arise from our perception of a combination of volatile ester compounds. Phenotypic variability in ester production was investigated using a segregating population from a ‘Royal Gala’ (RG; high ester production) × ‘Granny Smith’ (GS; low ester production) cross, as well as in transgenic RG plants in which expression of the alcohol acyl transferase 1 (AAT1) gene was reduced. In the RG × GS population, 46 quantitative trait loci (QTLs) for the production of esters and alcohols were identified on 15 linkage groups (LGs). The major QTL for 35 individual compounds was positioned on LG2 and co‐located with AAT1. Multiple AAT1 gene variants were identified in RG and GS, but only two (AAT1‐RGa and AAT1‐GSa) were functional. AAT1‐RGa and AAT1‐GSa were both highly expressed in the cortex and skin of ripe fruit, but AAT1 protein was observed mainly in the skin. Transgenic RG specifically reduced in AAT1 expression showed reduced levels of most key esters in ripe fruit. Differences in the ripe fruit aroma could be perceived by sensory analysis. The transgenic lines also showed altered ratios of biosynthetic precursor alcohols and aldehydes, and expression of a number of ester biosynthetic genes increased, presumably in response to the increased substrate pool. These results indicate that the AAT1 locus is critical for the biosynthesis of esters contributing to a ‘ripe apple’ flavour.     

Genetic influences on ‘environmental’ factors

Childhood environment, social environment and behavior, leisure time activities and life events have been hypothesized to contribute to individual differences in cognitive abilities and physical and emotional well‐being. These factors are often labeled ‘environmental’, suggesting they shape but not reflect individual differences in behavior. The aim of this study is to test the hypothesis that these factors are not randomly distributed across the population but reflect heritable individual differences. Self‐report data on Childhood Environment, Social Environment and Behavior, Leisure Time Activities and Life Events were obtained from 560 adult twins and siblings (mean age 47.11 years). Results clearly show considerable genetic influences on these factors with mean broad heritability of 0.49 (0.00–0.87). This suggests that what we think of as measures of ‘environment’ are better described as external factors that might be partly under genetic control. Understanding causes of individual differences in external factors may aid in clarifying the intricate nature between genetic and environmental influences on complex traits.     

Differential repair of excision gaps generated by transposable elements of the ‘Ac family’

Studies on transposable elements of the Ac family have led to different models for excision gap repair in either plants or Drosophila. Excision products generated by the plant transposable elements Ac and Tam3 imply a more or less straightforward ligation of broken ends; excision products of the Drosophila P element indicate the involvement of ‘double-strand break’ (DSB) repair. Recent findings that excision products of Ac and Tam3 can also contain traces of the element ends indicate, however, that DSB repair might be an alternative repair mechanism in plants. A functional DSB repair mechanism in plants can also be deduced from the observed rapid increases of Ac copy number during plant development and from the involvement of Ac in the generation of internal Ac deletions. On the other hand, alternative repair mechanisms may also be functional in Drosophila, because some of the ‘footprints’ generated upon P excision can be explained by a mechanism that has been postulated for excision gap repair in plants. It is concluded that plants and Drosophila can use similar repair mechanisms, but that the predominance of a certain repair mechanism is determined by the host.     

The rise of bilaterians: a few closing comments

Freeman (2009, 2010) argue that Ediacara “faunas” should be analysed using “promorphologies” of extant animals, that rangeomorphs and erniettomorphs are colonial animals, and that Vernanimalcula is a bilaterian animal. Recent studies of Ediacara fossils have provided multiple lines of evidence that these fossils represent a sample of phylogenetically diverse marine organisms that included crown-group animals as well as stem-group animals and non-animals. Thus, it is inappropriate to analyse Ediacara fossils using “promorphologies” of extant animals. The interpretation of rangeomorphs and erniettomorphs as colonial animals is inconsistent with the functional morphologies of these Ediacara fossils. Although Vernanimalcula is a fossil, the purported germ layers of Vernanimalcula are of diagenetic origin and there is no morphological evidence in support of its bilaterian affinity.     

Theraphosidae phylogeny: relationships of the ‘Ischnocolinae’ genera (Araneae,Mygalomorphae)

The mygalomorph spider subfamily ‘Ischnocolinae’ was originally established as a group based on the presence of divided tarsal scopula. Later, the divided condition of the scopula was considered the plesiomorphic state, which could not support the monophyly of ‘Ischnocolinae’. In Raven 1985, the subfamily was considered paraphyletic, pending a phylogenetic analysis to reinvestigate monophyletic groups. This study comprises such a phylogenetic analysis, based on morphological data, that includes representatives of all genera currently included in ‘Ischnocolinae’ as well as representatives of all other nine Theraphosidae subfamilies (Thrigmopoeinae, Ornithoctoninae, Eumenophorinae, Stromatopelminae, Harpactirinae, Selenogyrinae, Theraphosinae, Aviculariinae and Selenocosmiinae). The family Theraphosidae is considered monophyletic and expanded to include three additional genera previously considered as possible Barychelidae, namely Brachionopus (as Harpactirinae), Trichopelma and Reichlingia (as Ischnocolinae sensu stricto) while ‘Ischnocolinae’ as previously defined does not appear as monophyletic. However, two monophyletic groups were defined as subfamilies to include some former ‘Ischnocolinae’ representatives. The first group includes Acanthopelma rufescens, Trichopelma nitidum, Reichlingia annae, Ischnocolus spp., Holothele rondoni, Holothele culebrae and Holothele aff culebrae and is hereby named as Ischnocolinae (sensu stricto). The other subgroup comprises Sickius longibulbi, Holothele incei, Holothele aff incei, Guyruita spp., Schismatothele lineata, Hemiercus modestus, Holothele colonica and Holothele sp., together established as Schismatothelinae subfam. nov. Several genera included in former ‘Ischnocolinae’ appear as monophyletic (Catumiri, Oligoxystre, Heterothele, Nesiergus, Chaetopelma, Ischnocolus, Guyruita and Plesiophrictus). However, the genera Holothele, Schismatothele and Hemiercus deserve more attention in order to evaluate their intrarelationships and inclusion of species.     

‘Big data’ from shrinking pathogen populations

Falling costs for genome sequencing and genotyping mean that population genomic data sets are becoming commonplace for a wide variety of species. Once these data are used for the initial tasks of investigating population structure and demographic history, however, is there reason to go back for more? In this issue of Molecular Ecology, Nkhoma et al. (2013) explore the applications of longitudinal genomic diversity data for detecting changes in the prevalence and transmission of the Plasmodium falciparum malaria parasite in South‐East Asia. While this study finds several genetic signatures indicative of reduced disease transmission, other measures, such as short‐term effective population size, geographical population structure and heterozygosity, were not informative. These results indicate the potential contribution of genomic data to the surveillance of small, dynamic populations, whether they are at risk of extinction or targeted for elimination. The interpretation of such data will require close consideration of biological context, however, at both the species and the population level.     

Cleaner Wrasses Keep Track of the ‘When’ and ‘What’ in a Foraging Task1

In recent years evidence has accumulated that at least some animals can remember the ‘what’, ‘where’ and ‘when’ of personal experiences. Currently, evidence for such ability is taxonomically restricted to birds and mammals. Here, we demonstrate for the first time that cleaner wrasses Labroides dimidiatus are able to remember when they interacted with what after a single event. In nature, cleaners remove ectoparasites from other reef fishes, so‐called clients. Clients are depleted, non‐stationary food patches at the end of an interaction and replenished only after a delay. In our experiments, we presented twelve cleaners every 2.5 min, a choice between two of a total of four plates with different colours and patterns. One plate was always accessible but contained a non‐preferred food item while the other three contained a preferred food item, but allowed a next feeding event only after 5, 10 or 15 min. Thus, to maximise food intake, cleaners had to remember for each choice when they had last interacted with which plate. When confronted with two plates offering preferred food, cleaners showed an overall significant preference for the plate that allowed access during the trial. For six cleaners, the preference was significant. Also, on trials involving the always accessible plate, cleaners discriminated between trials in which they had to eat the non‐preferred food and trials on which they could eat the preferred food. In conclusion, cleaners are able to track the ‘when’ and ‘what’ (or possibly ‘who’) within a biologically meaningful time period.     

New early crown-group asteroids (Echinodermata; Triassic of Germany)

Migmaster angularis n. gen. n. sp. (Trichasteropsiidae, Forcipulatacea, Asteroidea) is described from the German Triassic Lower Muschelkalk of Anisian (Pelsonian) age, and new specimens ofTrichasteropsis bielertorum broaden understanding of this species. Phylogenetic events linking the stem groups, known only from Paleozoic rocks, with the crown group, known only from post-Paleozoic rocks, are problematic; the new fossils partially constrain phylogenetic changes associated with emergence of crown-group asteroids.      

The phylogeny of a reduced ‘sand goby’ group based on behavioural and life history characters

Phylogenetic analysis of 27 behavioural and life history traits for five Mediterranean sand goby species (Perciformes, Gobiidae) produced one tree with a consistency index (excluding uninformative characters) of 0.756. This tree agreed with previous molecular analyses in providing strong support for the monophyly of the sand gobies, indicating that Pomatoschistus and Knipowitschia are paraphyletic and helping to resolve the ambiguous position of Economidichthys pygmaeus, placing it as the basal member of the reduced data set. Although the tree was completely resolved, the branches above E. pygmaeus were only moderately supported in the bootstrap analysis. Overall, the behavioural data provide information that may eventually help clarify the speciation bursts within the Mediterranean sand goby clade as much as is possible. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 916–924.     

Potential effects of the invasive ‘killer shrimp’ (Dikerogammarus villosus) on macroinvertebrate assemblages and biomonitoring indices

1. Water quality monitoring data from 10 watercourses and laboratory mesocosm studies were used to assess the potential impacts of the crustacean amphipod invader Dikerogammarus villosus on resident macroinvertebrate assemblage structure in Central European fresh waters. 2. The presence of D. villosus was associated with a decline in the prevalence of many native species, pollution sensitive as well as pollution tolerant, and changes in biotic indices, despite the trends of improved water quality coinciding with the invasion period. A general increase in the prevalence of other invaders was also noted. The potential impacts of D. villosus were substratum dependent, differing between stone, concrete and sand‐dominated sites. 3. Mean Multimetric Macroinvertebrate Index Flanders (MMIF) values were marginally lower when D. villosus was present (P < 0.06), as opposed to when other amphipod species or no amphipods were present, despite the improved water quality. Mesocosm studies showed that several macroinvertebrate taxa were completely eliminated in treatments with D. villosus, oligochaete worms, Caenidae mayfly, chironomids and tipulids being particularly vulnerable to D. villosus predation. Biological Monitoring Working Party (BMWP) scores were lower in mesocosms with D. villosus as opposed to the native Gammarus pulex or no amphipods at all. 4. We predict that resident macroinvertebrate assemblages in both Central Europe and Britain will come under increasing pressure as D. villosus invasions progress. Consequently, macroinvertebrate biotic indices, such as the MMIF or BMWP, may need to be revised to account for changes in taxa sensitivities to water quality as well as increased predation and competition.     

Architectural complexity of marine crustacean burrows: unusual helical trace fossils from the Miocene of Mallorca,Spain

Gibert, J.M. de, Mas, G. & Ekdale, A.A. 2012: Architectural complexity of marine crustacean burrows: unusual helical trace fossils from the Miocene of Mallorca, Spain. Lethaia, Vol. 45, pp. 574–585. Unusual helical trace fossils occur in Tortonian shelfal calcarenites in the island of Mallorca. Their morphology may be either simple (ichnogenus Gyrolithes) or, more commonly, consist of two concentric helical burrows (ichnogenus Lapispira). They bear a very characteristic pelleted lining and are associated and probably connected to Thalassinoides and Ophiomorpha burrow systems very abundant in the same unit. These features allow the interpretation that the tracemaker was a thalassinidean shrimp. The complex and compound nature of these trace fossils is comparable to that seen in other modern and fossil crustacean burrow systems, and it reflects the behavioural plasticity of the architects. □Trace fossils, ichnology, ichnofabrics, crustacean burrows, Miocene, shallow marine, Lapispira.     

Developmental instability of vascular plants in contrasting microclimates at ‘Evolution Canyon’

Rocellaria dubia bores into subtidal rocks of karsted limestone in the Adriatic Sea and elsewhere. It also bores into the shells of various bivalve species. The mechanism of boring has hitherto been debated, but examination of occupied shells suggest that this is achieved by mechanical (the shell) abrasion and chemical etching using secretions produced from glands in the anterior mantle. Fast‐growing bivalves such as Ostrea edulis and Pinna nobilis carry heavy R. dubia burdens, and encapsulate the borer in secreted calluses. Slow‐growing bivalves such as the burrowing Venus verrucosa and Glycymeris violacescens carry low R. dubia burdens, are less able to encapsulate the borers, and probably incur enhanced mortalities as a result. Individuals of R. dubia removed from their limestone boreholes re‐secreted adventitious tubes around their siphons, probably from glands in the posterior mantle. The lifestyle of R. dubia is now better understood, and its ability to bore bivalve shells in particular suggests how the more advanced tropical gastrochaenids Cucurbitula and Eufistulana have evolved from initial (as juveniles) bivalve shell borers into occupants of adventitious crypts and tubes, respectively. It is further argued that the Gastrochaenidae show convergent similarities with the similar crypt‐ and tube‐building representatives of the Clavagelloidea. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 786–804.     

Complementary ‘bottom-up’ and ‘top-down’ approaches to basal ganglia function

Recently, two quite different approaches exemplifying ‘bottom-up’ and ‘top-down’ philosophies have shed new light on basal ganglia function. In vitro work using organotypic co-cultures has implicated the subthalamic nucleus (STN) and the external segment of the globus pallidus (GP_e) as pacemakers for low-frequency bursting that is reminiscent of the activity produced in Parkinsonian tremor. A circuit essential for avian song learning has been identified as part of the basal ganglia with surprisingly well conserved cellular details; investigation of this system may help to address general issues of basal ganglia function.     

ICHNOLOGICAL INSIGHTS INTO MITRATE PALAEOBIOLOGY

Abstract:  Mitrates are a controversial group of extinct deuterostomes; there is little agreement over their affinities, functional morphology or even the orientation of their upper and lower surfaces. Four slabs of slate from the Lower Devonian Hunsrück Slate (Bundenbach, Germany) are here described, showing trace fossils ( Vadichnites transversus igen. et isp. nov.) associated with the mitrate Rhenocystis latipedunculata . These new findings clearly demonstrate that the mitrate appendage was used in locomotion and that this movement took place appendage-first. Such a functional interpretation suggests that mitrates were oriented with the flat body surface upwards in life and argues against a phylogenetic position in the echinoderm crown-group.     

The ‘Hutchinsonian niche’ as an assemblage of demographic niches: implications for species geographic ranges

Hutchinson defined the ecological niche as a hypervolume shaped by the environmental conditions under which a species can ‘exist indefinitely’. Although several authors further discussed the need to adopt a demographic perspective of the ecological niche theory, very few have investigated the environmental requirements of different components of species’ life cycles (i.e. vital rates) in order to examine their internal niche structures. It therefore remains unclear how species’ demography, niches and distributions are interrelated. Using comprehensive demographic data for two well‐studied, short‐lived plants (Plantago coronopus, Clarkia xantiana), we show that the arrangement of species’ demographic niches reveals key features of their environmental niches and geographic distributions. In Plantago coronopus, opposing geographic trends in some individual vital rates, through different responses to environmental gradients (demographic compensation), stabilize population growth across the range. In Clarkia xantiana, a lack of demographic compensation underlies a gradient in population growth, which could translate in a directional geographic range shift. Overall, our results highlight that occurrence and performance niches cannot be assumed to be the same, and that studying their relationship is essential for a better understanding of species’ ecological niches. Finally, we argue for the value of considering the assemblage of species’ demographic niches when studying ecological systems, and predicting the dynamics of species geographical ranges.