Episodic chromosomal evolution in Planipapillus (Onychophora: Peripatopsidae): a phylogenetic approach to evolutionary dynamics and speciation

Planipapillus, a clade of onychophorans from southeastern Australia, exhibits substantial chromosomal variation. In the context of a robust phylogeny based on nuclear and mitochondrial sequence data, we evaluate models of chromosomal evolution and speciation that differ in the roles assigned to selection, mutation, and drift. Permutation tests suggest that all chromosome rearrangements in the clade have been centric fusions and, on the basis of parsimony and maximum-likelihood methods with independent estimates of branch lengths, we conclude that at least 31 centric fusions have been fixed in Planipapillus. A likelihood-ratio test approach, which is independent of our point estimates of ancestral states, rejects an evolutionary model in which the mutation rate is constant and centric fusions are effectively neutral. In contrast to the nucleotide sequence data, which are consistent with neutrality and rate constancy, centric fusions in Planipapillus are underdominant, spontaneous fusion rates vary among lineages, or both. We predict an inverse relationship between rates of chromosomal evolution and historical population size. Chromosomal evolution may play a role in speciation in Planipapillus, both by interactions between centric fusions with monobrachial homology and by the accumulation of multiple weakly underdominant fusions.     

Ultrastructure of mesoderm in embryos of Opisthopatus roseus (Onychophora, Peripatopsidae): revision of the "long germ band" hypothesis for Opisthopatus

In previous studies, an unusual pattern of development which resembles the "long germ band" development of some insects has been described in the onychophoran Opisthopatus cinctipes. This pattern has been proposed to be a characteristic of the genus Opisthopatus. To test this assumption, the ultrastructure of embryos of O. roseus, the sister species of O. cinctipes, was examined. Two kinds of paired, segmentally arranged coelomic cavities were found in the embryos studied: 1) dorsolateral coelomic cavities lined by extremely thin epithelia, and 2) ventral coelomic cavities situated within the anlagen of ventrolateral body appendages. Only the dorsolateral coelomic cavities can be considered "somites," since they occur earlier during embryogenesis. This is in contrast with the previous view that suggested a ventral position of "somites" in O. cinctipes. In addition, an anterior-to-posterior gradient occurs in the development of O. roseus. Based on our findings, we reevaluated the previous data on O. cinctipes. From this survey, no evidence in support of a "long germ band" hypothesis in Opisthopatus was found. Instead, the embryogenesis in representatives of Opisthopatus is more similar to that in other onychophorans than expected.     

The female genital system of Ooperipatellus decoratus (Onychophora, Peripatopsidae): an ultrastructural study

The female genital system of the oviparous peripatopsid Ooperipatellus decoratus consists of an ovary, oviducts equipped with receptacula seminis and additional pouches, uteri, and a vagina. It is examined using transmission and scanning electron microscopy. The ovary is made up of paired ovarian tubes united anteriorly and posteriorly and differentiated into a sterile dorsal part and a fertile ventral part with exogenous oocytes. Fertilization presumably occurs in the oviducts once the oocytes pass the receptaculum seminis. Although the receptacula seminis have been reported to occur in juvenile O. decoratus females only, the present study reveals that they are present in adult females as well. Their wall consists of a cuboidal epithelium covered with a thin collagen-muscle layer. The additional pouches are projections of the oviducts facing the receptacula seminis. They are distally closed to the haemocoel by a flattened epithelium and lack external muscle cells. A thin collagen layer is only found proximally. The uteri are characterized by a columnar epithelium with folded cell membranes allowing extension of the uteri, thus facilitating the passage of the large uterine eggs towards the vagina. Another dominating feature of the uteri is a distally increasing secretory production, which probably contributes to chorion development. Cilia occurring along the entire length of the uteri are considered to assist in the transport of eggs towards the vagina.     

An ultrastructural investigation of the hypocerebral organ of the adult Euperipatoides kanangrensis (Onychophora, Peripatopsidae)

The hypocerebral organs of Euperipatoides kanangrensis are a pair of spherical vesicles located ventral to the cerebral ganglia. They develop in the embryo from the most anterior pair of ventral organs, in the antennal segment. The wall of each hypocerebral organ is a dense epithelium of elongate cells with peripheral nuclei. The cytoplasm of the cells includes numerous mitochondria, Golgi bodies and microtubules. The small lumen, located eccentrically within the organ, contains concentrically layered electron-dense material resembling cuticle.Each hypocerebral organ is enclosed by a layer of extracellular matrix continuous with that surrounding the adjacent cerebral ganglion. There are no nerve connections between ganglion and organ, but cellular connections traverse the intervening matrix and could serve as a communication pathway. The ultrastructure of the hypocerebral organs indicates that they are glands.     

Variation in eggshell characteristics and in intrauterine egg retention between two oviparous clades of the lizard Lacerta vivipara: insight into the oviparity-viviparity continuum in squamates

The concept of the oviparity-viviparity continuum refers to the wide range in the length of intrauterine egg retention and, hence, in the stage of embryonic development at oviposition existing in squamates. The evolutionary process underlying this continuum may involve not only a lengthening of egg retention in utero, but also a marked reduction in the thickness of the eggshell. The idea that there may exist a negative correlation between the developmental stage reached by the embryo at oviposition and the eggshell thickness within squamates, although supported by the comparison of oviparous vs. viviparous species, has seldom been evaluated by comparing eggshell thickness of oviparous forms with different lengths of intrauterine egg retention. Eggs of two distinct oviparous clades of the lizard Lacerta vivipara were compared. The eggs laid by females from Slovenian and Italian populations have thicker eggshells, contain embryos on average less developed at the time of oviposition, and require a longer incubation period before hatching than the eggs laid by females from French oviparous populations. Our data and several other examples available from the literature support the idea that the lengthening of intrauterine retention of eggs and the shortening of the subsequent external incubation of eggs are associated with reduction in the thickness of the eggshell, at least in some lineages of oviparous squamates. The current hypotheses that may account for this correlation are presented and a few restrictions and refinements to those hypotheses are discussed. In particular, other changes, such as increased vascularization of the oviduct and of the extraembryonic membranes, may play the same role as the decrease of eggshell thickness in facilitating prolonged intrauterine egg retention in squamates. Future studies should also consider the hypothesis that the length of intrauterine retention might directly depend on the extent of maternal-fetal chemical communication through the eggshell barrier.     

Development and roles of vitelline cells in eggshell formation in Fasciola gigantica

Summary

In Fasciola gigantica, vitelline cells are the major contributors to the formation of the eggshell. The vitelline cells develop in vitelline follicles that are located in the posterior third of the adult parasite's body, in the areas lateral to the uterus and the testis. Mature vitelline cells are released and transported to the Mehlis' gland-ootype complex via a series of vitelline ducts. Based on ultrastructural features, the developing vitelline cells are classified into four stages: stem cell, protein-synthetic, carbohydrate-synthetic and mature cell stages. At the protein-synthetic stage, the eggshell globules are formed, whereas during the carbohydrate-synthetic stage glycogen particles and glycan vesicles are synthesized. The mature vitelline cells are detached from the nurse cells, and pass successively into the intrafollicular, interfollicular, longitudinal and transverse vitelline ducts, to be stored in the vitelline reservoir before being transported to the ootype via the median vitelline duct. At the same time, ova are transported from the ovary through the oviduct into the ootype lumen where each becomes surrounded by a number of vitelline cells. Vitelline cells secrete eggshell globules to surround a group of vitelline cells and an ovum in the ootype lumen, and these globules coalesce into the definitive eggshell. In the middle part of the uterus fertilization occurs, after which the eggshell is completely formed. Within the egg proper, vitelline cells break down, releasing glycogen and other products to nourish the developing embryo.     

Seminal receptacula in gravid and virgin female Peripatus (Macroperipatus) acacioi Marcus and Marcus (Onychophora,Peripatidae)

The ovoid seminal receptacula in Peripatus acacioi are located at the junctions of the short paired oviducts with the two horns of the uterus. Associated with each is a tubular funnel that opens into the haemocoel. In P. acacioi, spermatozoa may be stored in the seminal receptacula for several years (Campiglia and Walker '95, J. Morphol. 224:179–198). Observations of the structure of the seminal receptaculum using transmission electron microscopy (TEM) show that there are numerous tracheae within its wall indicating a good oxygen supply. The ultrastructure of the epithelium lining the seminal receptaculum indicates that these cells secrete the material that forms the luminal matrix that surrounds and provides nutrition for the stored spermatozoa. The ducts that interconnect the ovary, seminal receptaculum, funnel, and uterus are all packed with cilia. The structure of the seminal receptaculum and associated parts in the mature virgin female is identical to that of the gravid female, but the luminal matrix does not contain any spermatozoa. J. Morphol. 237:127–136, 1998. © 1998 Wiley-Liss, Inc.     

Biology of a thelyotokous biotype of Eretmocerus mundus (Hymenoptera: Aphelinidae) on Bemisia tabaci (Homoptera: Aleyrodidae)

Abstract Thelyotokous biotype of Eretmocerus mundus Mercet (Hymenoptera: Aphelinidae), a parasitoid of Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae), was recently recorded in northern Iran. Reproductive biology of this biotype was studied as part of an evaluation of its potential for biological control of B. tabaci. The parasitoid deposited more eggs under 2nd and 3rd nymphal instars than 1st or 4th instars. Adult females fed honey, with no access to whitefly nymphs, lived significantly longer (13.6 ± 4.7 d) than those given access to nymphs, but not fed honey (7.6 ± 2.21 d). Lifetime fecundity averaged 81.7 ± 26.9 female progeny per female parasitoid, ranging from 11–132. Daily fecundity, measured as the number of whitefly nymphs parasitized by per female each day for 10 d, averaged 18.06 ± 3.95 for the first 6 d of life, and then declined to < 11. Developmental time from oviposition to parasitoid emergence was significantly shorter in the 3rd instar of the host (15.9 ± 1.06 d) than in the 1st instar (18.7 ± 2.3 d), but not in the 2nd instar (16.4 ± 1.3 d).     

Characteristics of the sperm tail of the velvet worm, Euperipatoides leuckarti (Onychophora)

The onychophoran sperm tail contains several kinds of microtubulcs; probably more than that of any other animal group. There are thus a peripheral manchette consisting of many tightly spaced microtubules, a ring of nine 'peripheral singlets' and a central axoneme of the classical 9 + 2 type (nine doublets and two central singlets). The protofilament organization of these various microtubules was examined and compared to the structure and mode of formation of the peripheral singlets with that of its analogues in other animal groups. The onychophoran peripheral singlets were found to differ in two respects from those in insects: they are formed from the manchette rather than from the axonemal doublets and their transient connection to the axoneme is to the A-subtubules of the doublet rather than to the B-subtubules. The manchette microtubules as well as the peripheral singlets consist of 13 protoh'laments. The manchette may serve a mechanical function (to strengthen the unusually thick sperm tail) hut the role of the peripheral singlets remains unknown.     

The application of scanning electron microscopy to the systematics of the neotropical Peripatidae (Onychophora)

The taxonomy of the New World Peripatidae is poorly known both because of their rarity and difficulty in the determination of specimens. The distribution of papillae on the dorsal integument is an important taxonomic; character but is difficult to interpret under the light microscope. In this study the integument of a range of New World onychophoran species was studied using scanning electron microscopy. Twenty-one species were examined. Three species of Peripatopsidae were also studied for comparison. This technique revealed (a) the pattern of distribution of papillae on the integument, and (b) the morphology of the papillae. A pattern of three smaller papillae between two large was common to many species. Differences observed in some species were probably modifications of this basic pattern. There was interspecific variation in the number of scales in the apical and basal pieces of the papillae. The specimens examined fell into two main groups: (1) species with more than three scale ranks in the apical piece ( Oroperipatus and Peripatus ) and (2) species with three or fewer scale ranks ( Epiperipatus, Macroperipatus and others). The results thus support a distinction between Epiperipatus and Peripatus . However Macroperipatus is shown not to be a natural group. The relationships of some species which did not fit into this pattern are discussed. This paper shows scanning electron microscopy to be a potentially useful technique in a revision of the New World Peripatidae.