Past exam of the year 2 of the earth sciences (geology) course of the University of Oxford / 2023 / Paper 1 / 11

Section F: Evolution.

Gregor Mendel's first law is Mendelian segregation: the two alleles at a genetic locus in a diploid individual separate into different gametes, so each gamete receives one. In ordinary meiosis, separation of homologous chromosomes at meiosis I provides the chromosomal mechanism; their sister chromatids separate at meiosis II.

The second law is independent assortment: segregation at one genetic locus is independent of segregation at another when the loci are unlinked. Random orientation of different pairs of homologous chromosomes at meiosis I supplies the mechanism for genes on different chromosomes. Genetic linkage is an essential qualification: nearby loci on one chromosome do not generally assort independently, although sufficiently frequent genetic recombination can make their observed combinations approach independence. Genetic dominance concerns the phenotype of a heterozygote and is not one of these two segregation laws.

The biological species concept groups natural populations that actually or potentially interbreed and are reproductively isolated from other such groups. Its central criterion is reproductive isolation and gene flow, rather than visible similarity. It is hard to apply directly to fossils, organisms reproducing strictly by asexual reproduction, and geographically separated populations whose potential interbreeding cannot be tested.

The phenetic species concept groups organisms by overall similarity in measured phenotypic traits. A practical morphological species concept uses anatomical form as its main evidence; strict phenetic classification can combine many kinds of measured traits. This makes such approaches usable for preserved specimens and much of the fossil record, but the decision depends on which traits are measured and how similarities are weighted.

At least four difficulties arise. Cryptic species may be reproductively isolated while looking nearly identical, so similarity can lump separate lineages. Conversely, sexual dimorphism, juvenile/adult differences and phenotypic plasticity can make one interbreeding species look like several, causing over-splitting. Convergent evolution can make distantly related organisms resemble one another in ecologically selected traits. Finally, continuous geographic or temporal variation supplies no unique gap at which to place a boundary: character choice, sample size, preservation and similarity thresholds can alter the inferred species delimitation. These are distinct from the biological concept's practical difficulty of testing actual interbreeding.

Allopatric speciation begins when a geographic barrier divides a population and reduces gene flow. Independent mutation, genetic drift and divergent natural selection can then build reproductive isolation; if populations later meet, they remain distinct when isolation is sufficiently strong. Sister lineages of snapping shrimp in the genus Alpheus on opposite sides of the Isthmus of Panama illustrate the role of separated marine populations. Their divergences are not all synchronous with the final closure: genetic data indicate several separation histories, so one barrier date should not be assigned indiscriminately to every pair.^[ref]

Sympatric speciation develops without a geographic barrier separating the initial population. Divergent ecological selection combined with assortative mating, habitat-associated mating, different reproductive times or a chromosomal change can reduce gene flow within a shared area. The palms Howea forsteriana and Howea belmoreana on Lord Howe Island provide a documented candidate: different soil associations and substantially different flowering times contribute to reproductive isolation, and their sister relationship and timing support divergence on the island.^[ref]

The distinction concerns the geography of divergence, not whether present-day ranges overlap. Present coexistence alone cannot distinguish sympatric speciation from earlier allopatric speciation followed by range expansion. Parapatric speciation describes divergence between adjacent populations with limited exchange and provides an intermediate geographical category; the two main modes requested are the allopatric and sympatric endpoints.