Phenotypic evolution in a fossil gastropod species lineage: evidence for adaptive radiation?

Detecting speciation in the fossil record is a particularly challenging matter. Palaeontologists are usually confronted with poor preservation and limited knowledge on the palaeoenvironment. Even in the contrary case of adequate preservation and information, the linkage of pattern to process is often obscured by insufficient temporal resolution. Consequently, reliable documentations of speciation in fossils with discussions on underlying mechanisms are rare. Here we present a well-resolved pattern of morphological evolution in a fossil species lineage of the gastropod Melanopsis in the long-lived Lake Pannon. These developments are related to environmental changes, documented by isotope and stratigraphical data. After a long period of stasis, the ancestral species experiences a phenotypic change expressed as shift and expansion of the morphospace. The appearance of several new phenotypes along with changes in the environment is interpreted as adaptive radiation. Lake-level high stands affect distribution and availability of habitats and, as a result of varied functional demands on shell geometry, the distribution of phenotypes. The on-going divergence of the morphospace into two branches argues for increasing reproductive isolation, consistent with the model of ecological speciation. In the latest phase, however, progressively unstable conditions restrict availability of niches, allowing survival of one branch only.

Phenotypic evolution in a venerid bivalve species lineage from the late Middle Miocene Central Paratethys Sea: a multi-approach morphometric analysis

A morphometric analysis was performed for the late Middle Miocene bivalve species lineage of Polititapes tricuspis (Eichwald, 1829) (Veneridae: Tapetini). Specimens from various localities grouped into two stratigraphically successive biozones, i.e. the upper Ervilia Zone and the Sarmatimactra Zone, were investigated using a multi-method approach. A Generalized Procrustes Analysis was computed for fifteen landmarks, covering characteristics of the hinge, muscle scars, and pallial line. The shell outline was separately quantified by applying the Fast Fourier Transform, which redraws the outline by fitting in a combination of trigonometric curves. Shell size was calculated as centroid size from the landmark configuration. Shell thickness, as not covered by either analysis, was additionally measured at the centroid. The analyses showed significant phenotypic differentiation between specimens from the two biozones. The bivalves become distinctly larger and thicker over geological time and develop circular shells with stronger cardinal teeth and a deeper pallial sinus. Data on the paleoenvironmental changes in the late Middle Miocene Central Paratethys Sea suggest the phenotypic shifts to be functional adaptations. The typical habitats for Polititapes changed to extensive, very shallow shores exposed to high wave action and tidal activity. Caused by the growing need for higher mechanical stability, the bivalves produced larger and thicker shells with stronger cardinal teeth. The latter are additionally shifted towards the hinge center to compensate for the lacking lateral teeth and improve stability. The deepening pallial sinus is related to a deeper burrowing habit, which is considered to impede being washed out in the new high-energy settings.

Morphometric redefinition of the component chronospecies of the Globorotalia conoidea - G. inflata lineage in DSDP Hole 29-284

Phylo-zonations (or lineage-zonations) are based upon morphological changes within individual evolutionary lineages. These zonations, although potentially of use for stratigraphic subdivision and correlation, often suffer from a lack of quantitative exactness in the definitions of chronospecies. Thus exact reproducibility is hindered for stratigraphic determinations. The potential of morphometrically defined phylo-zonations is demonstrated on a temperate South Pacific Late Cenozoic lineage of planktonic foraminifera (Globorotalia conoidea through intermediate forms to Globorotalia inflata in DSDP Site 284) exhibiting phyletic gradualism. Our sampling interval is about 0.1 m.y. during the last 8 m.y. Changes in the number of chambers in the final whorl, test conicalness, percentage of keeled forms, and test roundness or inflatedness, are used to quantitatively define the following five chronospecies: G. conoidea (Late Miocene; 6.1->8.3 m.y.), G. conomiozea (latest Miocene ; 5.3-6.1 m.y.), G. puncticulata sphericomiozea (earliest Pliocene; 4.5-5.3 m.y.), G. puncticulata puncticulata (Early-Middle Pliocene; 2.9-4.5 m.y.), and G. inflata (Late Pliocene-Quaternary; 0-2.9 m.y.). This phylo-zonation is directly applicable to temperate cool subtropical Southern Hemisphere areas where the evolution took place (Kennett, 1967, 1973; Scott, 1979). It is still not known if the lineage occurs elsewhere; thus the applicability of the phylo-zonation over broader areas is still uncertain. Trends in general size and aperture shape seem to be climatically controlled, and thus may be only of local stratigraphic utility. The practical applications of morphometric phylo-zonation for stratigraphy is to a large extent dependent upon the amount of time and effort required to statistically define the trends. Experiments with large numbers of subsamples from this lineage demonstrate that accurate stratigraphic determinations are possible from measurements on only 15 specimens per sample, except for those very close to chronospecies boundaries.

Quantitative study of evolution in the Sphaeroidinella lineage in mid-Pleistocene sediment of ODP Hole 154-926A (Appendix A)

Morphological evolution in the late Neogene planktonic foraminifer Sphaeroidinella lineage involves a sudden increase of the percentage of specimens equipped with supplementary apertures (from <30% to >70%) in the mid-Pliocene (about 3.5 Ma). This evolutionary transition, marked by the first occurrence of specimens with large supplementary apertures in the lineage, is denoted the Sphaeroidinella event. Changes in the proportions of the supplementary apertures in the lineage were studied in 24 samples from ODP Hole 926A drilled in the equatorial Atlantic Ocean. In addition, detailed chronological models have been compiled for this section as well as for Pliocene sections from DSDP Holes 214, 502A, and 503B, where evolution in the lineage have been analyzed previously. Stratigraphic correlation of the studied sequences suggests that the Sphaeroidinella event took place at about 3.6 Ma in the eastern equatorial Pacific (Hole 503B) and at 3.5-3.6 Ma in the Caribbean (Hole 502A), while in the Atlantic Ocean (Hole 926A) and in the Indian Ocean (Hole 214) the event occurred after 3.5 Ma. The inferred diachrony of the mid-Pliocene Sphaeroidinella transition, which is considered to represent a prime example of punctuated anagenesis, suggests that this evolutionary modality may have an allopatric component. Its short duration (on average less than 50 kyr) and the detailed biochronology that could be established for this event qualifies it as a useful biostratigraphic tool in the low-latitude Pliocene oceans.