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  • Author or Editor: Menno Schilthuizen x
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The great diversity in genital shape and function across and within the animal phyla hamper the identification of specific evolutionary trends that stretch beyond the limits of the group under study. Asymmetry might be a trait in genital morphology that could play a unifying role in the evolutionary biology of genitalia. Here, I review the current knowledge on the taxonomic distribution, phylogenetic patterns, genetics, development, and ecology of asymmetric (chiral) genitalia. Asymmetric genitalia (male as well as female) have evolved from bilaterally symmetric ones (and sometimes vice versa), innumerous times in most animal taxa with internal fertilisation, and especially in Platyhelminthes, Arthropoda, Nematoda, and Chordata. In groups with asymmetric genitalia, chiral reversal (where species carry genitalia that are the mirror image of those in other, congeneric, species) is common, but antisymmetry (both mirror images present within a species) is rare. Although indications exist that, at least in insects, asymmetry evolves as a compensatory response to the evolution of male-dominant mating positions, many mysteries remain. Main questions are: (i) is genital asymmetry developmental-genetically linked with other (visceral, external) asymmetries? (ii) is genital asymmetry usually correlated with a change in mating position? (iii) is asymmetry more likely to evolve in response to cryptic female choice or sexually-antagonistic coevolution? (iv) why is antisymmetry so rare and how does chiral reversal evolve? Based on an overview of the taxonomic patterns, I advocate a research program that makes use of the simple, binary nature of left-right asymmetry to test hypotheses for its evolution with experimental and comparative methods. I also provide tables with full or summarised data on (a) genital asymmetry across all animal phyla with internal fertilisation; (b) genera with dextral as well as sinistral species; (c) species with dextral as well as sinistral individuals; (d) genera with symmetric as well as asymmetric species; (e) species with symmetric as well as asymmetric individuals.

Open Access
In: Animal Biology

Left-right asymmetry patterns in the body shapes of animals and plants have been a continuous source of interest among biologists. Recently, inroads have been made to developing a coherent research programme that makes use of the unique fact that chiral patterns may be studied (and generalities deduced) by comparisons across many unrelated groups, even across Kingdoms. The papers delivered at the symposium ‘Evolution of Chirality’ during the 2011 European congress of evolutionary biology (ESEB) provide examples of the various research programs that are currently developing within this field. The present paper provides a summary of the symposium, an introduction to this Special Issue of Contributions to Zoology, as well as suggestions for further collaboration among left-right asymmetry researchers.

Open Access
In: Contributions to Zoology

Since Solem’s provocative claim in the early 1980s that land snails in tropical forests are neither abundant nor diverse, at least 30 quantitative-ecological papers on tropical land snail communities have appeared. Jointly, these papers have shown that site diversity is, in fact, high in tropical forests; often more than 100 species have been recorded per site, which is somewhat more than normally found at sites in higher latitudes. At the same time, however, point diversities (which usually range between 10 and 30 species per quadrat) appear to be no different from the ones recorded for temperate localities, which suggests that the number of ways in which syntopic resource space can be subdivided among different land snail species has an upper limit that is no higher under tropical conditions. The available data do not allow much analysis of the ecological structuring processes of communities besides very coarse ones, e.g. the proportions of carnivores versus herbivores and Pulmonata versus non-pulmonates. Also, these first 30 years of research have shown that a number of serious methodological and conceptual issues need to be resolved for the field to move ahead; in particular whether empty shells from the forest floor may be used as a proxy for the contemporaneous communities. I make a number of suggestions for ways in which these obstacles may be removed. First, studies should be preceded by exploratory nested sampling in contiguous quadrats of increasing size, spanning several orders of magnitude. The shape of the triphasic species-area curve and nonlinear regression of the small-area end of the curve will help identify the quadrat and site areas that allow ecologically more meaningful studies. Second, researchers should be more aware of the trophic levels of species and restrict their analyses within guilds and within body size classes as much as possible. Testing species abundance distributions against ecologically explicit theoretical models may be a fruitful avenue for research. Finally, I argue that studies of this nature require species abundances that may only be found in tropical land snail communities that live on calcareous substrate, and therefore I suggest that malacologists aiming to understand community structure focus on limestone sites initially.

Open Access
In: Contributions to Zoology

The evolution of asymmetric genitalia is a common and recurrent phenomenon in a wide variety of insect taxa. However, little is understood about the evolution of left-right asymmetry in reproductive structures. Since a better knowledge of it could have an important impact on the study of genital evolution, in the present study we investigate the phylogenetic and evolutionary patterns of asymmetric male genitalia in Cyclocephalini. We use a Procrustes distance based method for quantifying asymmetry. Analysis of 119 species belonging to 14 genera revealed a diverse array of asymmetries with a strong indication that asymmetries are more strongly developed in the terminal part of the aedeagus. Further, we find that asymmetries have evolved repeatedly within this small taxon. Micro-CT scans, a technique not employed before in studies of genital asymmetry, are made of several symmetric and asymmetric species. This reveals unexpected asymmetric sclerotised structures inside the otherwise symmetric aedeagus of Cyclocephala amazona, which underlines that asymmetries are not restricted to the exterior of the male genitalia but are also found internally.

Open Access
In: Contributions to Zoology

The subtribe Anthroherponina form an iconic group of obligate cave beetles, typical representatives of the Dinaric subterranean fauna, which is considered to be the richest in the world. Phylogenetic studies within this subtribe are scarce and based only on morphological characters, which, due to troglomorphic convergence, are frequently unreliable. Moreover, morphological stasis and morphological polymorphism make classification of taxa difficult. To test if characters that have traditionally been accepted as informative for Anthroherponina classification are indeed reliable, we evaluated the monophyly of the most speciesrich genus of this subtribe - Anthroherpon Reitter, 1889. Our study, based on a molecular phylogenetic analysis of fragments of the 18S, 28S, and COI (both 5’ and 3’ end) loci revealed that the genus Anthroherpon as conventionally defined is polyphyletic. To resolve this polyphyly, we defined one new additional genus, Graciliella n. gen., for which we then examined the intrageneric diversity using molecular and morphometric approaches. Molecular phylogenetic analysis of two COI mitochondrial gene fragments revealed the presence of four species inside Graciliella n. gen., including two new species, which we here describe as G. kosovaci n. sp. and G. ozimeci n. sp. To analyze interspecific morphological differences within Graciliella we performed a discriminant analysis based on 40 linear morphometric measurements. The results showed that differences between species and subspecies inside Graciliella, however subtle they may seem, are measurable and reproducible. All species of the genus are briefly diagnosed, an identification key is proposed and a distribution map of all taxa of Graciliella is provided.

Open Access
In: Contributions to Zoology

The Southeast-Asian tree snail subgenus Amphidromus s. str. (Gastropoda Pulmonata: Camaenidae) is unusual among all gastropods for its genetic antisymmetry: populations consist of stable mixtures of individuals with clockwise (dextral) and counterclockwise (sinistral) coiling directions. Although previous studies in A. inversus suggest that this genetic dimorphism is maintained by sexual selection, it cannot be ruled out that environmental factors also play a role. Adult shell shapes in A. inversus are known to show subtle differences between both coiling morphs, and it is known that in snails in general, shell shape is under environmental selection, thus creating the possibility that micro-niche use of both coiling morphs differs. In this paper, we first confirm that hatchlings also differ in shell shape. We then proceed with field studies to compare dextral and sinistral juveniles and adults for (i) direction and rate of dispersal within the vegetation and (ii) micro-niche occupation. However, we failed to detect any difference in both ecological traits. In addition to earlier data that show that there is no clustering of morphs in the field and that both morphs suffer identical predation pressure, these new data do not provide any evidence for a role for environmental factors in maintaining the coil dimorphism in this species.

Open Access
In: Contributions to Zoology

In animals, cell polarity may initiate symmetry breaking very early in development, ultimately leading to whole-body asymmetry. Helical sperm cells, which occur in a variety of animal clades, are one class of cells that show clearly visible bilateral asymmetry. We used scanning-electron microscopy to study coiling direction in helical sperm cells in two groups of animals that have figured prominently in the sperm morphology literature, namely land snails, Stylommatophora (514 spermatozoa, from 27 individuals, belonging to 8 species and 4 families) and songbirds, Passeriformes (486 spermatozoa, from 26 individuals, belonging to 18 species and 8 families). We found that the snail sperm cells were consistently dextral (clockwise), whereas the bird sperm cells were consistently sinistral (counterclockwise). We discuss reasons why this apparent evolutionary conservatism of sperm cell chirality may or may not be related to whole-body asymmetry.

Open Access
In: Contributions to Zoology

Abstract

Here we present the first cases of birds using artificial plants as nest material. We report our findings for the common coot (Fulica atra) from Leiden, the Netherlands, in 2019. This is the first population of freshwater birds studied for its use of anthropogenic nest materials, and together with another report from the same year, the earliest case of an entire bird population with plastic in all nests. We also report the first artificial plants used as nesting material by birds, and discuss the implications of their usage as such.

Open Access
In: Behaviour

Abstract

Two new, extremely small land snail species, Angustopila coprologos Páll-Gergely, Jochum & Hunyadi n. sp. and Angustopila psammion Páll-Gergely, Vermeulen & Anker n. sp. are described from northern Vietnam and northern Laos, respectively. The former is characterized by a rough surface sculpture and bears tiny mud granules arranged in a pattern of radial lines on its shell surface. The latter species is the new global record-holder of the tiniest land snail title, with a shell width of 0.6–0.68 mm and a shell height of 0.46–0.57 mm. These measurements surpass the former records of Angustopila pallgergelyi and Acmella nana.

Open Access
In: Contributions to Zoology

Abstract

During the COVID-19 pandemic, Personal Protective Equipment (PPE) is massively used, resulting in a new wave of litter: protective face masks and gloves. Here we present the first case of a fish entrapped in a medical glove, encountered during a canal clean-up in Leiden, The Netherlands. We also report the first cases of birds using medical face masks as nesting material, which were also found in the Dutch canals. To place these new findings in context, we collected online reported interactions of animals with PPE litter, since the start of the pandemic. This resulted in the first overview of cases of entanglement, entrapment and ingestion of COVID-19 litter by animals and the use of it as nesting material. We signal COVID-19 litter as a new threat to animal life as the materials designed to keep us safe are actually harming animals around us. To understand the full scale of this problem, we welcome anyone to contribute to our overview by submitting their observations online at www.covidlitter.com. To further prevent PPE litter, it is recommended that, when possible, reusable alternatives are used.

Open Access
In: Animal Biology