Jonathan A. Todd

Middle Paleolithic shell beads in Israel and Algeria.

Perforated marine gastropod shells at the western Asian site of Skhul and the North African site of Oued Djebbana indicate the early use of beads by modern humans in these regions. The remoteness of these sites from the seashore and a comparison of the shells to natural shell assemblages indicate deliberate selection and transport by humans for symbolic use. Elemental and chemical analyses of sediment matrix adhered to one Nassarius gibbosulus from Skhul indicate that the shell bead comes from a layer containing 10 human fossils and dating to 100,000 to 135,000 years ago, about 25,000 years earlier than previous evidence for personal decoration by modern humans in South Africa.

Hopping Hotspots: Global Shifts in Marine Biodiversity

Hotspots of high species diversity are a prominent feature of modern global biodiversity patterns. Fossil and molecular evidence is starting to reveal the history of these hotspots. There have been at least three marine biodiversity hotspots during the past 50 million years. They have moved across almost half the globe, with their timing and locations coinciding with major tectonic events. The birth and death of successive hotspots highlights the link between environmental change and biodiversity patterns. The antiquity of the taxa in the modern Indo-Australian Archipelago hotspot emphasizes the role of pre-Pleistocene events in shaping modern diversity patterns.

Formation of the Isthmus of Panama

Independent evidence from rocks, fossils, and genes converge on a cohesive narrative of isthmus formation in the Pliocene. The formation of the Isthmus of Panama stands as one of the greatest natural events of the Cenozoic, driving profound biotic transformations on land and in the oceans. Some recent studies suggest that the Isthmus formed many millions of years earlier than the widely recognized age of approximately 3 million years ago (Ma), a result that if true would revolutionize our understanding of environmental, ecological, and evolutionary change across the Americas. To bring clarity to the question of when the Isthmus of Panama formed, we provide an exhaustive review and reanalysis of geological, paleontological, and molecular records. These independent lines of evidence converge upon a cohesive narrative of gradually emerging land and constricting seaways, with formation of the Isthmus of Panama sensu stricto around 2.8 Ma. The evidence used to support an older isthmus is inconclusive, and we caution against the uncritical acceptance of an isthmus before the Pliocene.

Environmental change preceded Caribbean extinction by 2 million years

Paleontologists typically treat major episodes of extinction as single and distinct events in which a major environmental perturbation results in a synchronous evolutionary response. Alternatively, the causes of biotic change may be multifaceted and extinction may lag behind the changes ultimately responsible because of nonlinear ecological dynamics. We examined these alternatives for the major episode of Caribbean extinction 2 million years ago (Ma). Isolation of the Caribbean from the Eastern Pacific by uplift of the Panamanian Isthmus was associated with synchronous changes in Caribbean near shore environments and community composition between 4.25 and 3.45 Ma. Seasonal fluctuations in Caribbean seawater temperature decreased 3-fold, carbonate deposition increased, and there was a striking, albeit patchy, shift in dominance of benthic ecosystems from heterotrophic mollusks to mixotrophic reef corals and calcareous algae. All of these changes correspond well with a simple model of decreased upwelling and collapse in planktonic productivity associated with the final stages of the closure of the isthmian barrier. However, extinction rates of mollusks and corals did not increase until 3–2 Ma and sharply peaked between 2 and 1 Ma, even though extinction overwhelmingly affected taxa commonly associated with high productivity. This time lag suggests that something other than environmental change per se was involved in extinction that does not occur as a single event. Understanding cause and effect will require more taxonomically refined analysis of the changing abundance and distribution patterns of different ecological guilds in the 2 million years leading up to the relatively sudden peak in extinction.

The ecology of extinction: molluscan feeding and faunal turnover in the Caribbean Neogene.

Molluscan faunal turnover in the Plio–Pleistocene of the tropical western Atlantic has been attributed to drops in temperature or primary productivity, but these competing hypotheses have not been assessed ecologically. To test these alternatives, we compiled data on changing molluscan life habits and trophic composition over 12 million years derived from 463 newly made collections from the southwestern Caribbean. Shelf ecosystems have altered markedly in trophic structure since the Late Pliocene. Predatory gastropods and suspension-feeding bivalves declined significantly in abundance, but not in diversity, and reef-dwellers became common. By contrast, all other ecological life habits remained remarkably stable. Food-web changes strongly support the hypothesis that declining regional nutrient supply had an increasing impact on regional macroecology, culminating in a faunal turnover.

Climate Change and Biosphere Response: Unlocking the Collections Vault

Natural history collections (NHCs) are an important source of the long-term data needed to understand how biota respond to ongoing anthropogenic climate change. These include taxon occurrence data for ecological modeling, as well as information that can be used to reconstruct mechanisms through which biota respond to changing climates. The full potential of NHCs for climate change research cannot be fully realized until high-quality data sets are conveniently accessible for research, but this requires that higher priority be placed on digitizing the holdings most useful for climate change research (e.g., whole-biota studies, time series, records of intensively sampled common taxa). Natural history collections must not neglect the proliferation of new information from efforts to understand how present-day ecosystems are responding to environmental change. These new directions require a strategic realignment for many NHC holders to complement their existing focus on taxonomy and systematics. To set these new priorities, we need strong partnerships between NHC holders and global change biologists.

Coral reef development drives molluscan diversity increase at local and regional scales in the late Neogene and Quaternary of the southwestern Caribbean

Abstract The late Neogene was a time of major environmental change in Tropical America. Global cooling and associated oceanographic reorganization and the onset and intensification of glaciation in the Northern Hemisphere during the past ten million years coincided with the uplift of the Central American isthmus and resulting changes in regional oceanographic conditions. Previous analyses of patterns of taxonomic turnover and the shifting abundances of major ecological guilds indicated that the regional shallow-water marine biota responded to these environmental changes through extinction and via a restructuring of local benthic food webs, but it is not clear whether this ecological response had an effect on the diversity of molluscan assemblages in the region. Changes in regional and local diversity are often used as proxies for similar ecological response to environmental change in large-scale paleontological studies, but a clear relationship between diversity and ecological function has rarely been demonstrated in marine systems dominated by mollusks. To explore this relationship, we have compiled a data set of the stratigraphic and environmental distribution of genera of mollusks in large new collections of fossil specimens from the late Neogene and Recent of the southwestern Caribbean. Analysis of a selection of ecological diversity measures indicates that within shelf depths, assemblages from deeper water (51–200 m) were more diverse than shallow-water (<50 m) assemblages in the Pliocene. Lower diversity for shallow-water assemblages is caused by increased dominance of a few superabundant taxa in each assemblage. This implies that studies of diversity of shelf benthos need to control for relatively fine scaled environmental conditions if they are to avoid interpreting artifacts of uneven sampling as true change of diversity. For shallow-water assemblages only, there was significant increase in local and regional diversity of bivalve assemblages after the late Pliocene. No parallel increase in gastropods could be detected, but this likely is because sample size was inadequate for documenting the diversity of gastropod assemblages following a steep post-Pliocene decline of average gastropod abundance. Both the increasing bivalve diversity and the decrease in average abundance of gastropod taxa correspond to an interval of increasing carbonate deposition and reef building in the region, and are likely a result of increased fine-scale habitat heterogeneity controlled by the local distribution of carbonate buildups. Each of these results demonstrates that documenting the ecological response of tropical marine ecosystems to regional environmental change requires a large volume of fine-scaled samples with detailed paleoenvironmental control. Such data sets are rarely available from the fossil record.

Amassing diversity in an ancient lake: Evolution of a morphologically diverse parthenogenetic gastropod assemblage in Lake Malawi

Exceptional ecological niche diversity, clear waters and unique divergent selection pressures have often been invoked to explain high morphological and genetic diversity of taxa within ancient lakes. However, it is possible that in some ancient lake taxa high diversity has arisen because these historically stable environments have allowed accumulation of lineages over evolutionary timescales, a process impossible in neighbouring aquatic habitats undergoing desiccation and reflooding. Here we examined the evolution of a unique morphologically diverse assemblage of thiarid gastropods belonging to the Melanoides polymorpha‘complex’ in Lake Malawi. Using mitochondrial DNA sequences, we found this Lake Malawi complex was not monophyletic, instead sharing common ancestry with Melanoides anomala and Melanoides mweruensis from the Congo Basin. Fossil calibrations of molecular divergence placed the origins of this complex to within the last 4 million years. Nuclear amplified fragment length polymorphism markers revealed sympatric M. polymorpha morphs to be strongly genetically differentiated lineages, and males were absent from our samples indicating that reproduction is predominantly parthenogenetic. These results imply the presence of Lake Malawi as a standing water body over the last million years or more has facilitated accumulation of clonal morphological diversity, a process that has not taken place in more transient freshwater habitats. As such, the historical stability of aquatic environments may have been critical in determining present spatial distributions of biodiversity.

Resistance of an invasive gastropod to an indigenous trematode parasite in Lake Malawi

Successful establishment and spread of biological invaders may be promoted by the absence of population-regulating enemies such as pathogens, parasites or predators. This may come about when introduced taxa are missing enemies from their native habitats, or through immunity to enemies within invaded habitats. Here we provide field evidence that trematode parasites are absent in a highly invasive morph of the gastropod Melanoides tuberculata in Lake Malawi, and that the invasive morph is resistant to indigenous trematodes that castrate and induce gigantism in native M. tuberculata. Since helminth infections can strongly influence host population abundances in other host-parasite systems, this enemy release may have provided an advantage to the invasive morph in terms of reproductive capacity and survivorship.

Environmental controls on shell-rich facies in tropical lacustrine rifts: A view from Lake Tanganyika's littoral

Abstract Lake Tanganyika, the worlds largest tropical rift lake, is unique among its counterparts in East Africa for the remarkable diversity of mollusk-rich sediments in its littoral zone. Molluscan shell beds are, however, a common feature of ancient lacustrine rift deposits and thus a better understanding of their spatial and temporal development is important. Targeted surveys across the littoral region of the Kigoma Basin reveal three surficial shell-rich facies that differ widely in depositional style and geometry. A unifying characteristic of these deposits is the volume of shells of Neothauma tanganyicense, a large, viviparous gastropod endemic to the lake. Reservoir-corrected radiocarbon dating indicates that Neothauma deposits in these surficial sediments are time averaged over at least the last ∼1600 calendar years BP. Preservation of fossil Neothauma shells in the littoral zone depends on both environmental conditions and on post-mortem shell modifications. Interaction between shells and mobile siliciclastic grains, facilitated by wave action and storms, represents a particularly destructive taphonomic process in the study area. Rank scoring of damage to Neothauma suggests that stromatolitic encrustations or early calcite coatings may help mitigate shell destruction caused by hydraulic fragmentation and abrasion. Persistence of Neothauma in littoral beds has important implications for the structuring of specialized communities of shallow-water benthos, as well as for improving analog models for hydrocarbon reservoirs in lacustrine carbonates.