Akiko Okusu

Evidence for a clade composed of molluscs with serially repeated structures: Monoplacophorans are related to chitons

Monoplacophorans are among the rarest members of the phylum Mollusca. Previously only known from fossils since the Cambrian, the first living monoplacophoran was discovered during the famous second Galathea deep-sea expedition. The anatomy of these molluscs shocked the zoological community for presenting serially repeated gills, nephridia, and eight sets of dorsoventral pedal retractor muscles. Seriality of organs in supposedly independent molluscan lineages, i.e., in chitons and the deep-sea living fossil monoplacophorans, was assumed to be a relict of ancestral molluscan segmentation and was commonly accepted to support a direct relationship with annelids. We were able to obtain one specimen of a monoplacophoran Antarctic deep-sea species for molecular study. The first molecular data on monoplacophorans, analyzed together with the largest data set of molluscs ever assembled, clearly illustrate that monoplacophorans and chitons form a clade. This “Serialia” concept may revolutionize molluscan systematics and may have important implications for metazoan evolution as it allows for new interpretations for primitive segmentation in molluscs.

Towards a phylogeny of chitons (Mollusca, Polyplacophora) based on combined analysis of five molecular loci

This study represents the first phylogenetic analysis of the molluscan class Polyplacophora using DNA sequence data. We employed DNA from a nuclear protein-coding gene (histone H3), two nuclear ribosomal genes (18S rRNA and the D3 expansion fragment of 28S rRNA), one mitochondrial protein-coding gene (cytochrome c oxidase subunit I), and one mitochondrial ribosomal gene (16S rRNA). A series of analyses were performed on independent and combined data sets. All these analyses were executed using direct optimization with parsimony as the optimality criterion, and analyses were repeated for nine combinations of parameters affecting indel and transversion/transition cost ratios. Maximum likelihood was also explored for the combined molecular data set, also using the direct optimization method, with a model equivalent to GTR + I + Γ that accommodates gaps. The results of all nine parameter sets for the combined parsimony analysis of all molecular data (as well as ribosomal data) and the maximum-likelihood analysis of all molecular data support monophyly of Polyplacophora. The resulting topologies mostly agree with a division of Polyplacophora into two major lineages: Lepidopleuridae and Chitonida (sensu Sirenko 1993). In our analyses the genus Callochiton is positioned as the sister group to Lepidopleuridae, and not as sister group to the remaining Chitonida (sensu BucklandNicks & Hodgson 2000), nor as the sister group to the remaining Chitonina (sensu Buckland-Nicks 1995). Chitonida (excluding Callochiton) is monophyletic, but conventional subgroupings of Chitonida are not supported. Acanthochitonina (sensu Sirenko 1993) is paraphyletic, or alternatively monophyletic, and is split into two clades, both with abanal gills only and cupules in the egg hull, but one has simple cupules whereas the other has more strongly hexagonal cupules. Sister to the Acanthochitonina clades is Chitonina, including taxa with adanal gills and a spiny egg hull. Schizochiton, the only genus with adanal gills that has an egg hull with cupules, is the sister-taxon to one of the Acanthochitonina clades plus Chitonina, or alternatively basal to Chitonina. Support values for either position are low, leaving this relationship unsettled. Our results refute several aspects of conventional classifications of chitons that are based primarily on shell characters, reinforcing the idea that chiton classification should be revised using additional characters.

Embryogenesis and development of Epimenia babai (Mollusca Neomeniomorpha).

Neomenioid aplacophorans (= Solenogastres) constitute one of the main lineages of molluscs. Developmental data of early embryogenesis and larval development of neomenioids are available for some species based on histological sections. I used other techniques to study the development of Epimenia babai Salvini-Plawen, 1997, and here I report new data on neomenioid development. The embryos of E. babai are lecithotrophic and cleavage is spiral, unequal, and holoblastic. Two polar lobes are formed, one at the first cleavage stage and one at the second cleavage stage. No evidence of external metameric iteration is visible through scanning electron microscopy or histology at any stage. A ciliated foot, a pedal pit, and aragonitic spicules develop from the definitive ectoderm. A spicule begins as a solid tip, continues to an open-ended hollow spicule, and finally becomes a closed-ended hollow spicule. The free-swimming trochophore larvae of E. babai have been considered unusual in lacking the characteristic neomenioid cellular test, an outer locomotory structure within which the entire definitive adult body develops. However, through the use of scanning electron and light microscopy, semithin sections, Hoechst nuclear staining, and programmed cell death staining to study the ontogeny and fate of the apical cells, I show that the entire pre-oral sphere (the apical cap) of the larvae is similar to the test of the other neomenioids. The results suggest that the test of the neomenioid larvae is an enlarged pre-oral sphere of a trochophore. The test morphologies of neomenioid larvae are compared to those of pericalymma larvae of protobranch bivalves, and the homology and evolution of molluscan larval tests is discussed.

Large population size predicts the distribution of asexuality in scale insects.

Understanding why some organisms reproduce by sexual reproduction while others can reproduce asexually remains an important unsolved problem in evolutionary biology. Simple demography suggests that asexuals should outcompete sexually reproducing organisms, because of their higher intrinsic rate of increase. However, the majority of multicellular organisms have sexual reproduction. The widely accepted explanation for this apparent contradiction is that asexual lineages have a higher extinction rate. A number of models have indicated that population size might play a crucial role in the evolution of asexuality. The strength of processes that lead to extinction of asexual species is reduced when population sizes get very large, so that the long‐term advantage of sexual over asexual reproduction may become negligible. Here, we use a comparative approach using scale insects (Coccoidea, Hemiptera) to show that asexuality is indeed more common in species with larger population density and geographic distribution and we also show that asexual species tend to be more polyphagous. We discuss the implication of our findings for previously observed patterns of asexuality in agricultural pests.

Phylogenetic analysis reveals positive correlations between adaptations to diverse hosts in a group of pathogen-like herbivores

A jack of all trades can be master of none—this intuitive idea underlies most theoretical models of host‐use evolution in plant‐feeding insects, yet empirical support for trade‐offs in performance on distinct host plants is weak. Trade‐offs may influence the long‐term evolution of host use while being difficult to detect in extant populations, but host‐use evolution may also be driven by adaptations for generalism. Here we used host‐use data from insect collection records to parameterize a phylogenetic model of host‐use evolution in armored scale insects, a large family of plant‐feeding insects with a simple, pathogen‐like life history. We found that a model incorporating positive correlations between evolutionary changes in host performance best fit the observed patterns of diaspidid presence and absence on nearly all focal host taxa, suggesting that adaptations to particular hosts also enhance performance on other hosts. In contrast to the widely invoked trade‐off model, we advocate a “toolbox” model of host‐use evolution in which armored scale insects accumulate a set of independent genetic tools, each of which is under selection for a single function but may be useful on multiple hosts.

Armored Scale Insects (Hemiptera: Diaspididae) of San Lorenzo National Park, Panama, with Descriptions of Two New Species

ABSTRACT Armored scale insects include invasive economic pests that have been widely studied in human-altered habitats but have received less attention in natural habitats. Although armored scale insects are nearly ubiquitous associates of woody plants, they generally go uncollected in general surveys because they are not susceptible to mass collecting techniques, such as fogging, beating, or trapping. San Lorenzo National Park in Panama was the subject of a recent high-profile effort to quantify the arthropod diversity in a tropical forest (Basset et al. 2012). Here, we contribute to understanding the biodiversity of this classic site by reporting the armored scale insect species we found there in August 2010. We found that, unlike other rainforest canopy taxa, the armored scale insect fauna is dominated by highly polyphagous cosmopolitan pests. However, we also found new species, and we describe two of them here: Furcaspis douglorum Okusu & Normark n. sp. and Hemiberlesia andradae Okusu & Normark n. sp. We provide amendments to the relevant keys, including a new key to New World species of Hemiberlesia Cockerell that lack perivulvar pores. In this context, we treat Abgrallaspis Balachowsky as asubjective synonym of Hemiberlesia, and we transfer its species to Hemiberlesia, except for the following assignments to other genera Affirmaspis flavida (De Lotto), n. comb., Aspidiotus furcillae Brain, rev. comb., Clavaspis perseae (Davidson), n. comb., Diaspidiotus fraxini (McKenzie), n. comb., and Davidsonaspis aguacatae (Evans, Watson, and Miller), n. comb. Davidsonaspis Normark, n. gen. We regard Abgrallaspis azadirachti Ojha and A narainus Dutta & Singh as synonyms of Aonidiella orientalis (Newstead), new synonymy.