Diversity

Most generalized biological definition

Sum total of variations in form, functioning, and behavior that
have accumulated in different lineages. Those variations
generally are adaptive to prevailing conditions or were once
adaptive to conditions that existed in the past.

Most commonly used definition

The number of species in a defined geographic area or in the
world

Diversity Indices - Sample-level measures of diversity

Richness [S]
S = number of species
* a measure strongly dependent on sample size (needs rarefying)

Shannon-Weaver Index [H] - measures both richness & evenness

H = - Σ Pi log Pi
Pi - a proportion of i-th species in a sample.
* For a given number of taxa H largest when all taxa equally abundant.
* relative abundance data needed for computation

Evenness (Equitability) Index [E]

E = H / Hmax
* Hmax is the maximum possible H at a given S and H is the actual
value computed for the analyzed sample.
* E varies from 0 to 1 (if all species are equally abundant E = 1).
* relative abundance data needed for computation
 Measures of Diversity

Taxic diversity - any diversity measures calculated using

number of higher taxa recorded in the fossil record in a
given time-interval

Relatively simple to obtain and easy to use

The problems:
(1) higher taxa are rarely monophyletic
(2) different taxa of the same taxonomic rank may be not
comparable (across groups or through time)
(3) without considering phylogeny, sampling biases
affecting diversity are more difficult to account for.

Nevertheless, an important & still successful method

Phylogenetic diversity - any diversity measures calculated

using number of lineages implied by phylogenetic trees
constructed from phena recorded in the fossil record.

Less prone to biases than the taxic approach

More difficult to apply on the larger scale (analysis is done
on species level).

Morphological disparity - any diversity measure based on

direct estimates of morphological variability in a given
group in a given time-interval.
Phylogeny & Taxic Diversity
Marine Phanerozoic Diversity
 Diversity vs. Turnover
• Diversity (standing diversity) – number of species
(lineages, higher taxa, or morphotypes) in a given time-
interval or given area

• Turnover (rate of change in diversity) –change in

proportion of unique species (lineages, higher taxa, or
morphotypes) through time or space

Diversity as a scale-dependent measure

Alpha diversity - Point-diversity (“Species Richness”)
Number of species within a single community or site. In
paleontological practice, number of species recorded in a
given fossil site.

Beta diversity
Turnover rate among communities. The amount of diversity
increased when two or more samples are combined.

Gamma diversity
Similar to beta diversity but measured on larger scale.
Turnover rate among regions or biogeographic provinces.