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This chapter reviews the current knowledge of variation in sperm morphology over several levels of biological organization: variation within males (both within and across ejaculates), among males, among populations, and among species, along with prevailing hypotheses addressing the adaptive significance of such variation. With regard to developmental mechanisms, three aspects of the physiology of sperm production serve to limit within-ejaculate variation in sperm phenotypes. First, the location of the testes and numerous aspects of testicular physiology of some taxa are clearly adaptations to maintain a homeostatic developmental environment for sperm. Second, developing spermatids may share cytoplasm. Third, sperm phenotypes are predominantly determined by testicular gene expression and hence the diploid genome of the male. Variation across ejaculates but within males can involve several traits including sperm numbers, overall semen quality and individual sperm quality. A special case of intramale variation in sperm form is found in species with sperm heteromorphism, in which different sperm forms are regularly produced by individuals. Differences among males in sperm morphology may derive from both genetic and environmental influences. Theories of condition-dependence basically posit that fitness-related traits are to a large extent dependent on an organism's underlying condition. Conclusions drawn from studies of sperm diversification between natural populations are reinforced by experimental evolution studies of sperm morphology in laboratory populations, as these studies address the evolvability of sperm traits and the nature of selection underlying sperm diversification. Furthermore, a discussion of evolutionary causes and consequences of sperm diversification, along with suggestions of fruitful areas for future exploration is presented.

Sperm morphological diversity

We conducted partitioned and combined Bayesian and parsimony phylogenetic analyses of corytophanid lizards (Iguania) using mtDNA, gross morphology, and sperm ultrastructure data sets. Bayesian and parsimony hypotheses showed little disagreement. The combined analysis, but not any of the partitioned ones, showed strong support for the monophyly of Corytophanidae and its three genera, Basiliscus, Corytophanes, and Laemanctus. Basiliscus is the sister taxon of a well-supported clade formed by Corytophanes and Laemanctus. The relationships of species within Basiliscus and Corytophanes received weak support, regardless of the method used. We defend those relationships as feasible and open to further testing. Data derived from the ultrastructure of spermatozoa are potentially a good source of characters for systematic inferences of Iguania and its major lineages. A Brooks Parsimony Analysis based on the geographic distributions of corytophanids and the phylogenetic tree obtained from the combined analysis suggested a Central American origin of the group, a recent colonization of northern South America, and the role of epeirogenic uplifts and the formation of lowlands during the late Tertiary in the differentiation of corytophanids.

Phylogenetic relationships of corytophanid lizards (Iguania, Squamata, Reptilia) based on partitioned and total evidence analyses of sperm morphology, gross morphology, and DNA data

Spermatogenesis, with emphasis on spermiogenesis, is described for the lizard, Tropidurus itambere, using light microscopy, phase contrast and epifluorescence, as well as scanning and transmission electron microscopy. Cellular differentiation involves events of chromatin condensation, nuclear elongation and the formation of structural complexes, such as the acrosomal and axonemal ones. Other new characteris- tics, exclusive for this species, include various aspects of the subacrosomal granule, the insertion of the pro- acrosomal vesicle and the development of these structures to participate in the acrosomal complex. Radial projections occur just above the nuclear shoulders, which have been recognized already from the beginning of cellular elongation. The development of the midpiece, the dense bodies, formation of the flagellum and elimi- nation of residual cytoplasm result in the final characterization of the mature spermatozoon. Comparisons between Tropiduridae and other lizard families are made.

Sperm ultrastructure and spermatogenesis in the lizard, Tropidurus itambere.

We describe the morphology of the spermatozoa of members of the iguanian lizard families Hoplocercidae and Opluridae for the first time. Their spermatozoa are quite similar and resemble the sperm of other iguanian lizards. Through a re-evaluation of sperm descriptions supplied in the literature we coded continuous and discrete characters through a combination of four methods: gap-weighting, step-matrix gap-weighting, reductive coding and composite coding. Parsimony analyses revealed that different methods produced highly incongruent outcomes. In view of the pros and cons of each method, we defend that the combination between composite coding and step-matrix gap-weighting are the most appropriate for our data. The cladogram derived from this analysis is totally resolved and comparatively well supported. We conclude that although the preferred cladogram was not strongly supported in some instances, sperm characters could be a good predictor of relationships among iguanian groups. We compare our hypothesis with preceding studies and we show that some previous groupings are in accordance with our results. Finally, we discuss the validity of those groupings in the light of methodological reasoning.

Sperm ultrastructure of hoplocercid and oplurid lizards (Sauropsida, Squamata, Iguania) and the phylogeny of Iguania

The body of ultrastructural data on spermatid characters during spermiogenesis continues to grow in reptiles, but is still relatively limited within the squamates. This study focuses on the ontogenic events of spermiogenesis within a viviparous and continually spermatogenic lizard, from high altitude in Mexico. Between the months of June and August, testicular tissues were collected from eight spermatogenically active bunchgrass lizards (Sceloporus bicanthalis) from Nevado de Toluca, Mexico. The testicular tissues were processed for transmission electron microscopy and analyzed to access the ultrastructural differences between spermatid generations during spermiogenesis. Interestingly, few differences exist between S. bicanthalis spermiogenesis when compared with what has been described for other saurian squamates. Degrading and coiling membrane structures similar to myelin figures were visible within the developing acrosome that are likely remnants from Golgi body vesicles. During spermiogenesis, an electron lucent area between the subacrosomal space and the acrosomal medulla was observed, which has been observed in other squamates but not accurately described. Thus, we elect to term this region the acrosomal lucent ridge. This study furthers the existing knowledge of spermatid development in squamates, which could be useful in future work on the reproductive systems in high altitude viviparous lizard species.

https://www.tandfonline.com/doi/pdf/10.4161/spmg.20410

Ontogenic development of spermatids during spermiogenesis in the high altitude bunchgrass lizard (Sceloporus bicanthalis)

The spermatozoa of Uta stansburiana and Urosaurus ornatus show the following squamate autapomorphies: a single perforatorium extending anteriorly from the apical tip of the paracrystalline subacrosomal cone; the presence of an epinuclear electron lucent region; intermitochondrial dense bodies; and the fibrous sheath extending into the midpiece. The acrosome vesicle is flattened and concentrically zoned apically; basally it overlies a subacrosomal cone which invests the nuclear rostrum. A stopper-like perforatorial base plate, rounded nuclear shoulders and a basal nuclear fossa are present. The proximal centriole contains a density within its centre for approximately one half its length and lies at approximately 80 degrees to the distal centriole. The two central singlets of the axoneme extend into the short distal centriole. A peripheral dense fibre is associated with each of the nine triplets of the distal centriole, and the fibre continues posteriorly with each of the nine doublets of the axoneme. A central fibre is associated with the two central singlets. All fibres are absent or vestigial at the level of the annulus. Mitochondria are short sinuous with a maximum of eight seen in transverse section. Uta and Urosaurus sperm differ from each other in their arrangement of intermitochondrial dense bodies in two ways: 1) longitudinally, Uta has five incomplete 'rings' of dense bodies, whereas Urosaurus has only four such rings; 2) in cross section, each individual 'ring' of Uta may contain up to four irregularly spaced dense bodies, whereas Urosaurus contains a maximum of only two dense bodies. The sperm of Uta and Urosaurus show strong similarities to those of the agamids and polychrotids. No spermatozoal autapomorphies for the Phrynosomatidae were found.

Morphology of the spermatozoa of the iguanian lizards Uta stansburiana and Urosaurus ornatus (Squamata, Phrynosomatidae).

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Morphology of the spermatozoa of the iguanian lizards Uta stansburiana and Urosaurus ornatus (Squamata, Phrynosomatidae).