Late postnaupliar development of the freshwater copepods Lovenula falicifera and Metadiaptomus colonialis (Calanoida: Diaptomidae) from South Africa

The life cycle of calanoid copepods consists of eggs hatching into nauplii (6 stages) which then moult into copepodids (5 stages), followed by the final moult into the adult female and male. The family Diaptomidae contains two subfamilies, Diaptominae and Paradiaptominae, with paradiaptomids almost exclusively consisting of African taxa. The copepodid stages III, IV and V were described for some freshwater diaptomine genera (i.e., Eudiaptomus Kiefer, 1932, Aglaodiaptomus Light, 1938, Skistodiaptomus Light, 1939, Leptodiaptomus Light, 1938, Megadiaptomus Kiefer, 1936 and Diaptomus Westwood, 1836). Copepods collected from Turfloop Dam, South Africa, with a plankton net were fixed and preserved in 70% ethanol. Calanoid copepods were studied under stereo- and light microscopes, using the wooden slide technique and features drawn. Examined specimens were identified as the copepodid stages of two African species, Lovenula falcifera (Lovén, 1845) and Metadiaptomus colonialis (van Douwe, 1914). Copepodids of the two species can be distinguished by their body size and the structure and size of the maxillipeds. The description and illustrations of three postnaupliar stages (CoIII, CoIV and CoV) are provided for both species. The identification of different stages is based on the number of urosomites, antennule development, the segmentation of legs 1–4, and the development of the fifth leg. These copepodids are compared with those of other described diaptomid genera.

The innovation of the final moult and the origin of insect metamorphosis

The three modes of insect postembryonic development are ametaboly, hemimetaboly and holometaboly, the latter being considered the only significant metamorphosis mode. However, the emergence of hemimetaboly, with the genuine innovation of the final moult, represents the origin of insect metamorphosis and a necessary step in the evolution of holometaboly. Hemimetaboly derives from ametaboly and might have appeared as a consequence of wing emergence in Pterygota, in the early Devonian. In extant insects, the final moult is mainly achieved through the degeneration of the prothoracic gland (PG), after the formation of the winged and reproductively competent adult stage. Metamorphosis, including the formation of the mature wings and the degeneration of the PG, is regulated by the MEKRE93 pathway, through which juvenile hormone precludes the adult morphogenesis by repressing the expression of transcription factor E93, which triggers this change. The MEKRE93 pathway appears conserved in extant metamorphosing insects, which suggest that this pathway was operative in the Pterygota last common ancestor. We propose that the final moult, and the consequent hemimetabolan metamorphosis, is a monophyletic innovation and that the role of E93 as a promoter of wing formation and the degeneration of the PG was mechanistically crucial for their emergence. This article is part of the theme issue ‘The evolution of complete metamorphosis’.

Copulation with immature females increases male fitness in cannibalistic widow spiders

Copulatory cannibalism of male ‘widow’ spiders (genus Latrodectus ) is a model example of the extreme effects of sexual selection, particularly in L. hasselti and L. geometricus where males typically facilitate cannibalism by females and mate only once. We show that these males can increase their reproductive success by copulating with final-instar, immature females after piercing the female's exoskeleton to access her newly developed sperm storage organs. Females retain sperm through their final moult and have similar fecundity to adult-mated females. This is an adaptive male tactic because immature mating increases insemination success relative to adult mating (which predicts higher paternity) and moreover, rarely ends in cannibalism, so males can mate again. Although successful only during a brief period before the female's final moult, males may employ this tactic when they associate with final-instar females in nature. Consistent with this, one-third of L. hasselti females collected as immatures in nature were already mated. Immature mating alters sexual selection on these otherwise monogynous males, and may explain male traits allowing facultative polygyny in Latrodectus . Since male cohabitation with immature females is common among invertebrates, immature mating may be a widespread, previously unrecognized mating tactic, particularly when unmated females are of high reproductive value.

The ontogeny and phylogeny of copepod antennules

Comparative analysis of the development of antennulary segmentation and setation patterns across six orders of copepods revealed numerous common features. These features are combined to produce a hypothetical general model for antennulary development in the Copepoda as a whole. In this model most compound segments result from the failure of expression of articulations separating ancestral segments. In adult males, however, compound segments either side of the neocopepodan geniculation are typically formed by secondary fusion at the last moult from CoV (stage 5). The array of segments distal to the articulation separating segments XX and XXI is highly conserved both in ontogeny and phylogeny: typically the distal segmentation of the adult female is already present in the CoI. A maximum of three setae is added to the distal array during the entire copepodid phase. This morphological conservatism is interpreted as evidence of the functional continuity of the distal setal array as a mechanosensory system providing early warning of approaching predators. Sexual dimorphism typically appears late in development; the male undergoing modifications especially at the final moult to sexual maturity. These modifications include the formation of the neocopepodan geniculation at the XX to XXI articulation and, in some orders, the formation of a proximal geniculation at the XV to XVI articulation. A proximal geniculation is reported here from the Calanoida for the first time. The geniculations allow the male to grasp the female during any mate guarding and during spermatophore transfer. Particular setae on segments either side of the neocopepodan geniculation are modified as basally fused spines in at least some representatives of the Calanoida, Misophrioida, Cyclopoida, Harpacticoida and Siphonostomatoida. The antennulary chemosensory system, comprising primarily the aesthetascs, is enhanced at the final moult in many male copepods. In planktonic copepods this enhancement may take the form of a doubling of the aesthetascs on almost every antennulary segment, as in the eucalanid calanoids, or of an increase in size of existing aesthetascs, as in the siphonostomatoid Pontoeciella , or of the transformation of possibly originally bimodal, seta–like elements into distally thin–walled, more aesthetasc–like elements, as in some calanoids, harpacticoids and poecilostomatoids. Enhancement of the chemosensory capacity of adult males appears to be linked with their mate–locating role. Copepods inhabiting the open–pelagic water column are more likely to exhibit enhancement of the chemosensory system than neritic or benthic forms. Enhancement may confer a greater sensitivity to chemosensory signals, such as pheromones produced by receptive females, which may retain their directional information at lower concentrations and, therefore, for longer periods, in oceanic waters than in more turbulent neritic waters. Aesthetascs appear to be more evolutionarily labile than other setation elements, apparently being lost and regained within well–defined lineages. Caution is urged in the use of aesthetasc patterns in phylogenetic analysis. The ontogenetic analyses suggest that the timing of expression of intersegmental articulations during development may in future provide the most informative characters for phylogenetic study, rather than either segment numbers or the patterns of fused or undivided segments.

The relationship between pre-flight reproduction and migratory urge in alatae of Aphis fabae Scopoli (Hemiptera: Aphididae)

Alatae of Aphis fabae Scop. which reproduced before their first flight (flyers) were compared in the laboratory with those which did not (migrants) in order to test whether this difference in behaviour could be taken as an expression of migratory urge. No difference in live weight or dry weight could be found between the two groups. Flyers, however had a higher wingloading than migrants after the final moult but not after pre-flight reproduction. Flyers also had a lower fat/dry weight ratio than migrants and appeared to have more embryos, including a greater number in an advanced state of development. The significance of these observations is discussed in relation to two hypotheses which could account for the different behaviour of migrants and flyers. The results support the hypothesis that migrants have an innately greater urge to take off than flyers, mediated by their crowding experience during development, and that this is likely to result in migratory as opposed to trivial flight.

Growth and moulting in nematodes: moulting and development of the hatched larva of Rotylenchulus reniformis

SummaryThe morphology of the post-hatch moulting and developmental sequence of an amphimictic population of Rotylenchulus reniformis has been examined in living and fixed material. Under suitable conditions of temperature and in a moist environment the 2nd-stage larva (L2) undergoes a series of 3 moults and developmental changes which result in the formation of males and immature females. After hatching and prior to the start of the first of these moults (the second moult), there is a pre-moult period, usually of 3–5 days duration. The morphology of the entire moulting and developmental sequences, from L2 to just after the final moult, has been followed in single living specimens of a parasitic nematode using differential interference contrast optics, and sections cut through different stages have been observed under the transmission electron microscope. These moulting sequences (at 24 °C) take place at similar times in developing males and females. The second moult takes place on the second day after commencement of moulting, the third on the third or fourth days and the fourth and final moult on the sixth or seventh days, followed by further development over several days to give rise to the adult male and the immature female. The ultrastructure associated with some of the more obvious of these developmental changes is described and includes the morphology of the head region and cuticles of L2, L4, adult males and immature females, the oesophageal glands of the immature female and the copulatory spicules and spermatozoa of the male.

Water content, weight change, and activity of apterous and alate virginoparous Acyrthosiphon pisum (Harris) (Homoptera: Aphididae)

Changes in weight, water content, and activity of alate and apterous virginoparous Acyrthosiphon pisum (Harris) were observed during development from third instar larvae to mature adults. Apterous aphids gained weight steadily until the 4th day of adulthood, and showed no dramatic changes in activity during this period. The live weights of alate aphids decreased during the 24 h immediately following the final moult, but increased gradually after this lime. The onset of weight loss occurred before the active nonfeeding period during which dispersal would normally lake place. Recommencement of weight gain followed the resumption of feeding. After the adult moult, the water content of alatae expressed as a percentage of live weight decreased for 24 h. whereas that of apterae remained constant. During the next 24 h, the water content of alatae increased to a level slightly below that of apterae. Thereafter, both morphs maintained a constant water content until about the 5th day, when a slight increase was evident. Eighty-five percent of the weight loss of adult alatae is attributable to water loss. It is suggested that dehydration of newly moulted adult alatae is an adaptation to facilitate dispersive flight.

Studies on the Malayan forest rat filaria, BreinUa booliati (Filarioidea: Onchocercidae): Course of development in rat host

The development of Breinlia booliati is described in its natural host, Rattus sabanus and in an inbred strain of laboratory albino rat. The growth of the parasite is similar in both the rat hosts. The third moult occurs between six—eight days and the final moult between 24–28 days. Larvae were recovered initially from the skin and carcass. After five weeks, developing stages were seen only in the thoracic and abdominal cavities, the site of development of the adult worms. Worms became sexually mature by 11–12 weeks and there was considerable growth in length of the female worms after this stage.

The effects of host diet on Thelastoma attenuatum (Nematoda: Thelastomatidae) populations in cockroaches

Experiments were constructed to assess the ellects or changes in the diet of Periplaneta americana on Thelastoma attenuatum. Synthetic diets depressed parasitaemia although apparently adequate for the host. However, adult worms were supported best by high carbohydrate levels although protein was necessary for the final moult. Little effect was observed on the distribution of worms within the htndgut although excess roughage appeared to result in a slight backward migration. Egg output per worm remained fairly constant but rose slightly under conditions of extreme stress. Dietary changes affected the size of adult female worms differentially but had little apparent effect on food reserves. Possible reasons for the changes are discussed