SUMMARY1. With few exceptions, copepods dominate over other crustacean and non-crustacean invertebrate groups in ground water. They have colonised a vast array of habitats in continental ground waters, where they are represented by over 1000 species in six orders: Platycopioida, Misophrioida, Calanoida, Cyclopoida, Harpacticoida, Gelyelloida. However, members of only the last four orders entered genuine fresh ground water. 2. Stygobiotic copepods show a wide range of morphological and physiological adaptations to different groundwater habitats. They frequently exhibit simplifications in body plans, including reductions in appendage morphology, which is regarded as a result of paedomorphic heterochronic events. 3. Copepod distributions at small spatial scales are most strongly affected by habitat type and heterogeneity, with sediment grain size and availability of organic matter being important habitat characteristics. Large-scale spatial distributions (biogeographical) are mainly related to past geological, climatic and geographical processes which occurred over medium to long time scales. 4. Such processes have affected colonisation patterns and diversification of copepods in ground water, leading to a number of phylogenetic and distributional relicts and a high degree of endemism at different taxonomic levels. This is reflected in the composition of groundwater copepod communities characterised by distantly related species in the phylogenetic tree. 5. Copepods dominate the species richness of groundwater fauna in all regions and on all continents where more than cursory surveys have been carried out, i.e. in Europe, North and Latin America as well as in Australia. 6. Species-specific microhabitat preferences, high proportions of local endemics, high proportions of phylogenetic and distributional relicts, and higher-level taxonomic diversity are all factors suggesting that copepods are a useful indicator group of overall species richness for defining conservation priorities in ground water.
Mate location and recognition are essentially asymmetrical processes in the reproductive biology of calanoid copepods with the active partner (the male) locating and catching the largely passive partner (the female). This behavioural asymmetry has led to the evolution of sexual dimorphism in copepods, playing many pivotal roles during the various successive phases of copulatory and post-copulatory behaviour. Sexually dimorphic appendages and structures are engaged in (1) mate recognition by the male; (2) capture of the female by the male; (3) transfer and attachment of a spermatophore to the female by the male; (4) removal of discharged spermatophore(s) by the female; and (5) fertilization and release of the eggs by the female. In many male calanoids, the antennulary chemosensory system is enhanced at the final moult and this enhancement appears to be strongly linked to their mate-locating role, i.e. detection of sex pheromones released by the female. It can be extreme in calanoids inhabiting oceanic waters, taking the form of a doubling in the number of aesthetascs on almost every segment, and is less expressed in forms residing in turbulent, neritic waters. Mate recognition is a process where chemoreception and mechanoreception presumably work in conjunction. The less elaborate male chemosensory system in the Centropagoidea is counterbalanced by females playing a more active role in generating hydromechanical cues. This is reflected in females in the shape of the posterior prosomal margin, the complexity of urosomal morphology and the size of the caudal setae. Visual mate recognition may be important in the Pontellidae, which typically show sexual dimorphism in eye design. The most distinctive sexual dimorphism is the atrophy of the mouthparts of non-feeding males, illustrating how copepod detection systems can be shifted to a new modality at the final moult. In the next phase, the male captures the female using the geniculate antennule and/or other appendages. Three types of antennulary geniculations are recognized, and their detailed morphology suggests that they have originated independently. Grasping efficiency can be enhanced by the development of supplemental hinges. The scanty data on capture mechanisms in males lacking geniculate antennules are reviewed. It is suggested that the loss of the antennulary geniculation in many non-centropagoidean calanoids has evolved in response to increasing predator pressure imposed on pairs in amplexus. Spermatophore transfer and placement are generally accomplished by the modified leg 5 of the male. In some males, leg 5 consists of both a chelate grasping leg and a spermatophore-transferring leg, whereas in others, only the latter is developed. Tufts of fine setules/spinules and/or sclerotized elements on the terminal portion of the leg are involved in the transfer and attachment of the spermatophore. The configuration of gonopores, copulatory pores and their connecting ducts in the female genital double-somite is diversified in the early calanoid offshoots such as Ari...
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 modi¢cations especially at the ¢nal moult to sexual maturity. These modi¢cations 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 ¢rst 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 modi¢ed 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 ¢nal 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-de¢ned lineag...
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