CHAPTER SUMMARY Here we outline the meaning of the term alternative reproductive tactics, or ARTs, and discuss why the existence of ARTs is so widespread in animals. We ask what we need to know to understand the evolution of ARTs and the importance of general principles such as frequency dependence, density dependence, and condition dependence, and what we need to know about proximate mechanisms involved in the regulation of ARTs to comprehend evolutionary patterns. We discuss current issues in the study of ARTs and list 12 questions that we think need particular attention. Throughout we shall provide representative examples of ARTs in animals to illustrate the ubiquitous nature of this phenomenon. WHAT IS THE MEANING OF ALTERNATIVE REPRODUCTIVE TACTIC ? Alternative The concept of ARTs refers to alternative ways to obtain fertilizations in both males and females. In its most common use, this term refers to traits selected to maximize fitness in two or more alternative ways in the context of intraspecific and intrasexual reproductive competition. In general, alternative phenotypes are characterized by a discontinuous distribution of traits evolved towards the same functional end. Examples include size dimorphism, color polymorphism, dimorphic morphological structures involved in the monopolization of resources or mates, and various behavioral alternatives such as territoriality vs. floating, monopolization vs. scramble competition, or investment in primary access to a resource vs. social parasitism. Individuals allocate resources to either one or the other (mutually exclusive) way of achieving the same functional end using evolved decision-making rules (Brockmann 2001).

The evolution of alternative reproductive tactics: concepts and questions

A review of the largely confused Amblyomma maculatum Koch, 1844 tick group of the subgenus Anastosiella Santos Dias, 1963 (A. neumanni Ribaga, 1902, A. maculatum, A. parvitarsum Neumann, 1901, A. tigrinum Koch, 1844 and A. triste Koch, 1844) is presented together with a discussion of the diagnostic characters used for the determination of adults, nymphs and, to a lesser extent, larvae. A key for this tick group is produced, including the description of the larva of A. parvitarsum, 1901. Sequences of 16S rDNA are obtained and compared with other Amblyomma spp., including two other species currently in Anastosiella but in the ovaletick group, A. ovale Koch, 1844 and A. aureolatum (Pallas, 1772). According to the morphology and the rDNA sequences, the maculatum group is reduced to A. maculatum (Neotropical-Nearctic), A. tigrinum (Neotropical) and A. triste (Neotropical) A. neumanni and A. parvitarsum are excluded from the subgenus. The distribution is sympatric in northern South America from where A. maculatumreaches the southern Nearctic and the range of A. tigrinum extends to the southern Neotropics. These species have been found on several domestic and wild vertebrates. A. triste and A. tigrinum have been also found on man. Their role as vectors of pathogens deserves further investigation.

The Amblyomma maculatum Koch, 1844 (Acari: Ixodidae: Amblyomminae) tick group: diagnostic characters, description of the larva of A. parvitarsum Neumann, 1901, 16S rDNA sequences, distribution and hosts.

The number of de novo mutations in the germline can be expected to increase with age in males, therefore females might decrease mutation load in their progeny by avoiding mating with older males. Here, I propose that female polyandry can be more effective in decreasing the risk of genetic disorders in progeny than pre-copulatory mate choice, particularly if sperm competitiveness declines more steeply with age than other traits affecting chances of males to mate. If faster ageing of spermatogenic tissue causes older males to transfer inadequate numbers of functional sperm, polyandry would also benefit females directly.

Male age, germline mutations and the benefits of polyandry

Host specificity of Neotropical hard ticks (Acari: Ixodidae) was analyzed by using the number of hosts species for each tick species and the index of host specificity S(TD)*, which integrates phylogenetic and ecological information. The analyses were based on 4172 records of hard ticks collected from wild and domestic tetrapods. Most tick species included in this study were associated with three to 20 host species. No tick species has been associated either with a single species or with a single genus of host. It was found that the number of host species is sensitive to sampling effort, but not the S(TD)*. The most frequent values of S(TD)* were between 2.5 and 3.5, which shows that the host species more frequently used by Neotropical hard tick species belong to different families or different orders. Immature stages tend to use a broader taxonomic range of hosts than adults, and the interpretation of both measures of host specificity used in this study led to the conclusion that the impact of non-endemic hosts does not alter the patterns of host specificity in Neotropical hard ticks. The index S(TD)* showed that a high proportion of tick species has phylogenetically unrelated species as principal hosts. The conclusion reached in this work indicates that strict host specificity is not common among Neotropical hard ticks and suggests that the influence of tick ecology and evolution of habitat specificity, tick generation time, phenology, time spent off the host and the type of life-cycle could be more important than hosts species.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/C5C513467D6C94B36786F523F36CE7B6/S0007485312000557a.pdf/div-class-title-a-meta-analysis-of-host-specificity-in-neotropical-hard-ticks-acari-ixodidae-div.pdf

A meta-analysis of host specificity in Neotropical hard ticks (Acari: Ixodidae).

Sperm competition is a well-recognised agent in the evolution of sperm and ejaculate structure, as well as variation in female quality. Models of the evolution of ejaculate expenditure predict that male body condition, female fecundity and the risk and intensity of sperm competition may be the ultimate factors shaping optimal ejaculate size. We investigated sperm allocation in Austropotamobius italicus, a freshwater crayfish exhibiting a coercive mating system and external fertilisation, in relation to male and female traits and copulation behaviour under laboratory conditions. We found that mating males were sensitive to female size and produced larger ejaculates when mating with larger females, which were more fecund in terms of number of eggs produced. We found no evidence for female egg production being sperm-limited, as the number of eggs was not dependent on male sperm expenditure. Copulation duration and number of ejaculations reliably predicted the amount of sperm transferred, and both these behavioural measures positively covaried with female body size. These results indicate that male freshwater crayfish can modulate their sperm expenditure in accordance with cues that indicate female fecundity. In addition, a novel finding that emerged from this study is the decrease in sperm expenditure with male body size, which may either suggest that large, old male crayfish are better able than small males to economise sperm at a given mating to perform multiple matings during a reproductive season, or that they experience senescence of their reproductive performance.

https://www.jstor.org/stable/info/25063805

Sperm allocation in relation to male traits, female size, and copulation behaviour in freshwater crayfish species

Two basic male morphs occur in several species of the family Acaridae: heteromorphic fighters, possessing a thickened and sharply terminated third pair of legs, and homeomorphic males with unmodified legs. We compared major life-history traits of the two morphs in the bulb mite, Rhizoglyphus robini. We found no significant differences in development time or virility, but homeomorphic males lived 23% longer than heteromorphs. We discuss the possibility that the trade-off between longevity and adaptation for fighting maintains genetic variation for the male morph in the studied species.

Iwona Bogacz

Papers

Comparison of life-history traits of the two male morphs of the bulb mite, Rhizoglyphus robini.