Preface to the Princeton Landmarks in Biology Edition vii Preface xi Symbols Used xiii 1. The Importance of Islands 3 2. Area and Number of Speicies 8 3. Further Explanations of the Area-Diversity Pattern 19 4. The Strategy of Colonization 68 5. Invasibility and the Variable Niche 94 6. Stepping Stones and Biotic Exchange 123 7. Evolutionary Changes Following Colonization 145 8. Prospect 181 Glossary 185 References 193 Index 201

The Theory of Island Biogeography

The diversity of life is one of the most striking aspects of our planet; hence knowing how many species inhabit Earth is among the most fundamental questions in science. Yet the answer to this question remains enigmatic, as efforts to sample the world's biodiversity to date have been limited and thus have precluded direct quantification of global species richness, and because indirect estimates rely on assumptions that have proven highly controversial. Here we show that the higher taxonomic classification of species (i.e., the assignment of species to phylum, class, order, family, and genus) follows a consistent and predictable pattern from which the total number of species in a taxonomic group can be estimated. This approach was validated against well-known taxa, and when applied to all domains of life, it predicts ∼8.7 million (±1.3 million SE) eukaryotic species globally, of which ∼2.2 million (±0.18 million SE) are marine. In spite of 250 years of taxonomic classification and over 1.2 million species already catalogued in a central database, our results suggest that some 86% of existing species on Earth and 91% of species in the ocean still await description. Renewed interest in further exploration and taxonomy is required if this significant gap in our knowledge of life on Earth is to be closed.

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How Many Species Are There on Earth and in the Ocean

It is clear that the above can lead to confusion when scientists of different countries are trying to communicate with each other. Another example is the burrowing rodent called a gopher found throughout the western United States. In the southeastern United States the term gopher refers to a burrowing turtle very similar to the desert tortoise found in the American southwest. One final example; two North American mammals known as the elk and the caribou are known in Europe as the reindeer and the elk. We never sing “Rudolph the Red-nosed elk”! Confused? This was the reason for creating an internationally recognized system of naming organisms. To avoid confusion, living organisms are assigned a scientific name based on Latin or Latinized words. The English sparrow is Passer domesticus or Passer domesticus (italics or underlining these two names is the official written representation of a scientific name). Using a uniform naming system allows scientists from all over the world to recognize exactly which life form a scientist is referring to. The naming process is called the binomial system of nomenclature. Passer is comparable to a surname and is called the genus, while domesticus is the specific or species name (like your given name) of the English sparrow. Now scientists can give all sparrow-like birds the genus Passer but the species name will vary. All similar genera (plural for genus) can be grouped into another, “higher” category (see below). Study the following for a more through understanding of taxonomy. Taxonomy Analogy Kingdom: Animalia Country

Diversity of life

https://www.cell.com/current-biology/pdf/S0960-9822(12)01138-4.pdf

The magnitude of global marine species diversity

Wolbachia are intracellular bacteria that manipulate the reproduction of their arthropod hosts in remarkable ways. They are predominantly transmitted vertically from mother to offspring but also occasionally horizontally between species. In doing so, they infect a huge range of arthropod species worldwide. Recently, a statistical analysis estimated the infection frequency of Wolbachia among arthropod hosts to be 66%. At the same time, the authors of this analysis highlighted some weaknesses of the underlying data and concluded that in order to improve the estimate, a larger number of individuals per species should be assayed and species be chosen more randomly. Here we apply the statistical approach to a more appropriate data set from a recent survey that tested both a broad range of species and a sufficient number of individuals per species. Indeed, we find a substantially different infection frequency: We now estimate the proportion of Wolbachia-infected species to be around 40% which is lower than the previous estimate but still points to a surprisingly high number of arthropods harboring the bacteria. Notwithstanding this difference, we confirm the previous result that, within a given species, typically most or only a few individuals are infected. Moreover, we extend our analysis to include several reproductive parasites other than Wolbachia that were also screened for in the aforementioned empirical survey. For these symbionts we find a large variation in estimated infection frequencies and corroborate the finding that Wolbachia are the most abundant endosymbionts among arthropod species.

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Still a Host of Hosts for Wolbachia : Analysis of Recent Data Suggests That 40% of Terrestrial Arthropod Species Are Infected

There is a bewildering range of estimates for the number of arthropods on Earth. Several measures are based on extrapolation from species specialized to tropical rain forest, each using specific assumptions and justifications. These approaches have not provided any sound measure of uncertainty associated with richness estimates. We present two models that account for parameter uncertainty by replacing point estimates with probability distributions. The models predict medians of 3.7 million and 2.5 million tropical arthropod species globally, with 90% confidence intervals of [2.0, 7.4] million and [1.1, 5.4] million, respectively. Estimates of 30 million or greater are predicted to have <0.00001 probability. Sensitivity analyses identified uncertainty in the proportion of canopy arthropod species that are beetles as the most influential parameter, although uncertainties associated with three other parameters were also important. Using the median estimates suggests that in spite of 250 years of taxonomy and around 855,000 species of arthropods already described, approximately 70% await description.

/pdf/quantifying-uncertainty-in-estimation-of-tropical-arthropod-41k89zj709.pdf

Quantifying uncertainty in estimation of tropical arthropod species richness.

There remains great uncertainty about how much tropical forest canopies contribute to global species richness estimates and the relative specialization of insect species to vertical zones. To investigate these issues, we conducted a four-year sampling program in lowland tropical rainforest in North Queensland, Australia. Beetles were sampled using a trap that combines Malaise and flight interception trap (FIT) functions. Pairs of this trap, one on the ground and a second suspended 15-20 m above in the canopy were located at five sites, spaced 50 m or more apart. These traps produced 29986 beetles of 1473 species and 77 families. There were similar numbers of individuals (canopy 14473; ground 15513) and species (canopy 1158; ground 895) in each stratum, but significantly more rare species in the canopy (canopy 509; ground 283). Seventy two percent of the species (excluding rare species) were found in both strata. Using IndVal, we found 24 and 27% of the abundant species (n>or=20 individuals) to be specialized to the canopy and the ground strata, respectively, and equivalent analyses at the family level showed figures of 30 and 22%, respectively. These results show that the canopy and the ground strata both provide important contributions to rainforest biodiversity.

/pdf/beetle-assemblages-from-an-australian-tropical-rainforest-zvi265cayp.pdf

Beetle assemblages from an Australian tropical rainforest show that the canopy and the ground strata contribute equally to biodiversity

The vertical stratification of insect species assemblages inhabiting tropical rainforests is well established but few have examined whether these patterns are reflected in vertical stratification of body size or feeding guilds. We used Malaise and Flight Interception Traps to sample beetle assemblages from five locations, at both canopy and ground zones of a tropical lowland rainforest site near Cape Tribulation, Australia. Beetles from 4 years of sampling were sorted to Family and morphospecies, and allocated to one of five feeding guilds. Within feeding guilds the number of species and individuals, from canopy- and ground-caught traps were compared. The body lengths of species were measure and compared within feeding guilds and families. Herbivores was the dominant guild but was not the majority of all species or individuals. Most beetle species (69%) were less than 5 mm in length and the mean size of canopy-caught species was greater than that for ground-caught species. This was probably due to slightly more species of plant feeders (herbivores and xylophages) present in the canopy, which were significantly larger than saprophages, fungivores and predators. Among feeding guilds, there were few overall canopy–ground differences. These results contrast with species composition results presented elsewhere where strong differences between the canopy and the ground were evident. We suggest that our guild groupings may have been too coarse to detect fine-scale differences and that resource partitioning may have also masked faunal stratification. We propose that fine-scale differences in resources between the canopy and the ground, together with strong microclimate gradients, are likely to be important in structuring the vertical stratification of insect assemblages at the level of species, but not with respect to functional groups.

Vertical stratification of feeding guilds and body size in beetle assemblages from an Australian tropical rainforest

How much do site age, habitat structure and spatial isolation influence the restoration of rainforest beetle species assemblages?

One of the least understood aspects of insect diversity in tropical rain forests is the temporal structuring, or seasonality, of communities. We collected 29,986 beetles
of 1473 species over a 4-yr period (45 monthly samples), with the aim to document the temporal dynamics of a trophically diverse beetle assemblage from lowland
tropical rain forest at Cape Tribulation, Australia. Malaise and flight interception traps were used to sample adult beetles at five locations at both ground and canopy
levels. Beetles were caught throughout the year, but individual species were patchy in their temporal distribution, with the 124 more abundant species on average being
present only 56 percent of the time. Climatic variables (precipitation, temperature, and solar radiation) were poorly correlated with adult beetle abundance, possibly
because: (1) seasonality of total beetle abundance was slight; (2) the peak activity period (September–November) did not correspond to any climatic maxima or
minima; or (3) responses were nonlinear owing to the existence of thresholds or developmental time-lags. Our results do not concur with the majority of tropical insect
seasonality studies suggesting a wet season peak of insect activity, perhaps because there is no uniform pattern of insect seasonally for the humid tropics. Herbivores
showed low seasonality and individual species’ peaks were less temporally aggregated compared to nonherbivores. Canopy-caught and larger beetles (> 5 mm) showed
greater seasonality and peaked later in the year compared to smaller or ground-caught beetles. Thus seasonality of adult beetles varied according to the traits of feeding
ecology, body size, and habitat strata

Peter S. Grimbacher

Papers

Quantifying uncertainty in estimation of tropical arthropod species richness.

Beetle assemblages from an Australian tropical rainforest show that the canopy and the ground strata contribute equally to biodiversity

Vertical stratification of feeding guilds and body size in beetle assemblages from an Australian tropical rainforest

How much do site age, habitat structure and spatial isolation influence the restoration of rainforest beetle species assemblages?

Seasonality of a Diverse Beetle Assemblage Inhabiting Lowland Tropical Rain Forest in Australia