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Commonness and rarity in species distribution
Commonness and rarity regards the patterns of relative abundance of species
disturbed across geographic space. According to Rabinowitz’s definition,
commonness and rarity should be defined based distribution range, habitat specialty
and population size, thus only species with large range, low habitat specialty and large
population size are regarded as common, and the other seven scenarios should be
various extent of rare. Of particular importance, what is common and rare depends on
the spatial scales under which species distribution is examined. The concerns of
species commonness and rarity is apparent from conservation standpoints: extinction
rate is directly related to degree of rarity, and having species exhibiting high rates of
extinction relative to others will lead to rare species. Moreover, how does
commonness-rarity pattern vary among taxonomic groups, geographic location, and
local species richness is important to understand community structure and biodiversity
patterns.
The main body of literatures are focused on assessing commonness and rarity
patterns, and investigate process and mechanisms explaining those patterns. Few
studies involved both goals, while majority of literatures especially before 2000
concentrated on the pattern assessment. I found there are several questions in
research regarding species commonness and rarity of importance: 1) is commonness
common? 2) does species distribution range and abundance correlate? 3) what is the
pattern of local abundance for rare species; 4) what processes account for being
common and rare? The last question gives rise to a rich body of research seeking for
mechanisms and processes that can explain species commonness and rarity.
Phylogenetic structures of common and rare species combined with species traits
(Philips et al, 2010), genetic variation between common and rare species (Cole, 2003),
congeneric species comparison (Münzbergova, 2005) generated and tested hypotheses
that why species become common and rare. While phylogenetic and population
genetic information is increasing accessible, how evolutionary history contributed to
commonness and rarity pattern is still poorly known.
[1] Schoener, T.W. (1987) Geographical distribution of rarity. Oecologia, 87, 161–
173.
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A dataset on bird species across Australia was used to analyse commonness and rarity
patterns. The occurrence data (frequency of being detected as presence) were
collected from 10,000km2 quadrats over the country which resulted in atlas. The
author aimed to examine whether the Australia birds tend to show suffusive or
diffusive rarity (see definition in summary), and what is their geographic distribution
range. Different quadrat size was used to show how spatial scales affect range size
distribution among species (e.g. more species showed restricted range when using
small quadrats and less tend to show restricted range using large quadrats). The major
discovers were 1) distributions of range size are strikingly skewed toward small
number of quadrats (most of species have restricted range) no matter which quadrat
size is used although large size showed less degree of this skewness; 2) diffusive
rarity (rare in some quadrat but not rare in others); 3) majority of species are common
(at least not rare) within their occurrence range; 4) range size is negatively correlated
with abundance, which did not support the hypothesis that wide-distributed species
have higher abundance (many taxonomic groups did not show any relationship
though). The study explicitly measured rarity which could realistically reflect the
patterns in natural communities, and involved datasets over wide geographic range,
thus provided a good example of assessing species rarity. Unfortunately there are not
very many studies similar in term of spatial scales following this research, probably
due to limitation of data availability, i.e. quite few datasets cover distribution rage, as
well as abundance, for a reasonable size of taxonomic groups.
[2] Pritt, J.J. & Frimpong, E.A. (2009) Quantitative determination of rarity of
freshwater fishes and implications for imperiled-species designations. Conserv.
Biol. 24, 1249–1258.
This study assembled datasets collected by REMAP and NAWQA programs resulted
in over 1300 stream sites and over 700 freshwater and anadromous species across the
North American. The objectives were to establish criteria for rarity on basic of
species range, habitat speciality and local population size, and to assess the
conservation status regarding their rarity. They quantified species range based on
distribution range by latitude and longitude, habitat speciality based on published
traits information (e.g. stream type, current and substrate conditions), and population
size based on assemblage datasets (abundance). They discovered that among 772,
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334 and 388 species were classified as respective range-extent common and rare
species, and among 678 species with available habitat use information, 369 and 309
species showed broad and narrow habitat uses, and over 30% species were rare
regarding population size. For 399 species of all the three type of information is
available, most abundant category is large distribution range, broad habitat use and
large population size followed different degrees of rarity in term of habitat use and
population size. This study provided a framework to access species commonness
rarity and could serve as tool to assess species imperilment status. However, the study
did not manage to avoid the problem derived from sampling bias, meaning
information for common (wide distributed and high abundance) species are more
available which probably explain why this category contributed large proportion
among the species to which the analyses are restricted. This also leads me to think
Rabinowitz’s definition is difficult to apply even with relatively extensive datasets,
and Schoener’s diffusive and suffusive rarity seems more practical.
[3] Pitman, N.C.A. et al. (2001) Dominance and distribution of tree species in upper
Amazonian Terra Firme forests. Ecology, 82, 2101–2117.
The study was conducted for tree species at two forests at the westernmost margin of
Amazon basin (Manu and Yasunĭ forest), with each area surveyed through respective
9 and 15 quadrats of approximately 1 hectare. The objectives were to study patterns
of commonness and rarity of the trophic tree species at local, landscape and regional
scales, in order detect whether commonness and rarity is associated with traits and
phylogenetic patterns (which family tends to have more common or rare species). In
both the Manu and Yasunĭ inventories, 150 species (out of ~500 species) had
landscape-scale densities of >1 individual/ha, and regarded as common, and majority
of rare species contributed minority of biomass and basal area. Common species
showed taxonomic pattern whereas rarity did not: most families had consistently
higher number of common species at both sites, whereas families showing higher
numbers of rare species did not show consistent trends (occurrence of rarity among
lineages appear to be random). Most species traits or attributes were not correlated
with landscape-scale abundance while they did find that monoecious species were
more abundant in these two forests compared with species with other reproductive
systems. The study tacked the questions whether certain lineages get dominant
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because of their life-history and morphological traits, and is among the few studies
that looked high diversity system, for which one may expect different commonness
and rarity patterns. One might question the use of threshold (>1 individual per
hectare) to define commonness, but seems did not affect their conclusion that rarity is
common in this study system. A limitation should be noticed that they did not
consider species distribution range which could be a source of errors because species
may appear rare at certain proportion of distribution range but not others.
[4] Cucherousset, J. et al. (2008) How do biodiversity patterns of river animals
emerge from the distributions of common and rare species? Biol. Conserv., 141,
2984–92.
This type of research asks the question how well the species richness is presented by
common and rare species using aquatic taxa (fishes, and macroinvertebrates) over
large drainage basin area in River Garonne, French. Among all the sites taxa (within
fishes, and different order of aquatic insects) were firstly ranked based on their
abundance/occurrence, then species were added and treated as sub-assemblage in
sequence of common to rare and rare to common, thus the correlation between subsets
of the taxa and overall species richness could be plotted as common versus rare
species were gradually added. Apart from plotting against adding species, the
relationship between correlation and cumulative information presented by species was
also examined, which was the cumulative sum across species of the binomial variance
∑pi (1 – pi), where pi = proportion of study area occupied by the ith species. The
correlation for common-to-rare species approached correlation coefficient = 1 more
rapidly than rare-to-common species, indicating the common species gave a closer
approximation of overall species richness pattern. Although similar results were also
reported from plants and small mammals, exception was seen from rare species in
fish, Ephemeroptera and Plecoptera when increasing ∑pi (1 – pi) (not species number)
along the common-to-rare and rare-to-common sequences. This result suggested that
rarer species showed a stronger affinity for high richness areas than do the commoner
species, and thus whether common or rare species better present richness also depends
on local species richness. This study demonstrated the peculiarity of aquatic taxa in
commonness and rarity questions, also pointed out common species showed closer
approximation of total taxa richness pattern probably only when the local diversity is
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low. There are quite a few analyses using same methods, and provide some
information on regional community diversity, however, the pattern revealed by this
methods are more like sampling artefact rather than the nature of assemblage
structures, also whether common or rare taxa are more preventative of total richness is
not relevant in make conservation decision.
[5] Arita, H.T. (1993). Rarity in Neotropical bats: correlations with phylogeny, diet,
and body mass. Ecol. Appl., 3, 506-517.
In order to examine rarity Neotropical bat species in relation to phylogenetic
constrains, diet and body mass, variations in distribution range and local abundance in
150 bat species were compared among taxonomic and trophic groups, and correlated
with maximum body mass. The body mass was not correlated with relative
abundance, but positively related with distribution range which was consistent from
the patterns observed in other taxanomic groups for which dispersal ability is thought
to be important shaping local community. Rarity was more frequently observed in
carnivores and insectivores species, and less occurred among species feeding on plant
parts. It was also found there was one family contain more rare species than others.
It was helpful to treat abundance and distribution range differently (did correlation
with them separately), because the scales and processes shaping abundance and range
are different. Because most proportion of variations in abundance was from amonggenus variation, and variations in distribution range from species, and the author
suggested this results indicated a phylogenetic constrains upon among-species
variations. However, the analytical approaches used in correlating relative abundance
and distribution range with phylogeny seems could not lead them to this conclusion.
[6] Mehranvar, L. & Jackson, D.A. (2001) History and taxonomy: their roles in the
core-satellite hypothesis. Oecologia, 127, 131–142.
Landbridge and oceanic islands, due to connectivity with mainland and historical
origin, should have different species distributions. Taxa with different dispersal
ability will occur on islands with different probability. The objective of this section is
to determine whether differences among taxa in their dispersal abilities match
particular patterns in their distributions (i.e. bimodal: toward either common or rare
with less intermediate abundance classes; Core: skewed toward more common
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species; satellite: skewed toward rare species; uniform distributions: none of the three
above patterns). The study compiled results of 108 case studies that surveyed island
habitat, and taxa include birds, mammals, amphibian and reptiles, arthropods, fish and
plants. They found more bimodal and core patterns in landbridge then oceanic island,
probably because of the fact that island seldom shared an ancestral biota (i.e.
opportunistic dispersal and colonization was major component in assemblages) that
limited the potential for having core species. Apparent variation on distribution
patterns comparing landbridge and oceanic island were revealed from different
taxonomic groups indicating an important role of dispersal ability. The result
suggested the species distribution pattern depended on the island history connected to
the mainland as well as species dispersal ability. The study explicitly tested the role
of dispersal in forming species abundance distribution patterns. There is a lack of
hypothesis testing studies (using either meta-analyses or field experiments), seems
that some studies that speculated on some mechanisms actually described the patterns
without a prior expectation of the possible causal mechanism.
[7] Hessen, D.O. and Walseng, B. (2008) The rarity concept and commonness of
rarity in freshwater zooplankton. Freshwat. Biol. , 53, 2026–2035.
Dataset on freshwater crustacean zooplankton across over 2000 Norwegian lakes were
used to assess and investigate potential causes of species commonness and rarity
patterns. Occurrence abundance were mainly used, i.e. the proportion of lakes (local
occurrence) and ecoregions (regional occurrence) species were reported as presence.
Based on Rabionowitz’s criteria, most of species were rare by distribution range: 65%
species were recorded from 10% area only 6 species were recorded more than 50%
localities. The major pattern found in this study was common species were common
by all the three criteria and vice versa. They suggested commonness and rarity was
inherent property, for some species they either have limited range because of life
cycle strategies or on the edge of distribution; many rare species showed high
dispersal and colonization ability, leading the author to ascribe their rarity to
competitive exclusion that occurred in the past. The study breakdown the pattern into
regionally wide distributed and restricted groups as well as into ecoregions, thus
displayed hierarchical spatial levels under which rarity and commonness was
examined. Yet, one of their conclusion is not convincing because they did not
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conduct analyses on species traits (except biomass), but speculated that species
distribution are not related to dispersal or colonization ability.
[8] Tales, E., Keith, P. & Oberdorff, T. (2004) Density-range size relationships in
French riverine fishes. Oecologia, 138, 360–370.
In order to test the principal biological mechanisms already proposed (i.e. the niche
breadth hypothesis and the resource availability hypothesis), the study examined data
for 738 reference sites covering a period of 13 years of survey (1985-1998) cross
French rivers. Species range size and local abundance was calculated and habitat
requirement, biological traits and physiologic traits were assembled from literatures.
They examined the relation between the local density of species and the size of the
geographic range for French stream fishes. Body size and other life history traits, as
well as habitat availability were correlated with species range size and local
abundance. As for most other taxonomic groups, a positive interspecific relationship
between abundance and range size was found. Their result also support resource
availability instead of niche breath hypothesis because species utilising resources
(habitats) that are marginal (less available) tend to appear at low density and to have
narrow distribution whereas species utilising widespread habitats tend to be more
abundant and more widely distributed. Only body size explains significant variation
around the relationship, being negatively correlated with local density and positively
correlated with range size. The study offered an alternative mechanism that is
especially important for aquatic species: habitat availability and distribution also
regulate species commonness and rarity.
[9] Cornwell, W.K. & Ackerly, D.D. (2010) A link between plant traits and
abundance: evidence from coastal California woody plants. J. Ecol., 98, 814–
821.
There is mounting evidence that plant communities in different biomes showed
trait-based presence/absence pattern, while species commonness and rarity could
be consequence of traits and stochastic events. In order to test whether species
relative abundance is correlated with species traits, and how this correlation vary
among different scales, a study was conducted at Jasper Ridge in California, US,
and 44 20mX20m plots were surveyed for woody tree species. Abundance was
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measured as individual and biomass, and eleven functional traits (e.g. specified
leaf area: SLA, nitrogen per leaf mass Nmass) were quantified. There are four
traits out of eleven, SLA, wood density, max height, and % lumen area, exhibited
effects upon species commonness and rarity at plot scale, demonstrating
influences of species traits upon their relative abundance, but not in landscape
scale. Also wood density showed opposite relationships between arid and wet
area indicating affinity between traits and abundance also depends on abiotic
environments. This study serves as a piece of evidence favouring trait-based,
non-random assembly rule against neutral assembly rule, and more important
whether trait-based process determine assemblage structures are scale dependent.
[10] Phillips, R.D. et al. (2010) Orchid biogeography and factors associated with
rarity in a biodiversity hotspot, the Southwest Australian Floristic Region. J.
Biogeogr., 38, 487–501.
The study is design to investigate the relative importance of pollination strategies,
edaphic habitat and site of mycorrhizal infection orchid taxon rarity in Southwest
Australian Floristic Region. They used geographic analyses to quantify rarity based
on the distribution data: the distributions of 407 orchid species were mapped at the
quarter-degree scale. The greatest number of rare taxa (defined as either low
abundance or small distribution) occurred in areas of high taxon richness and
naturally fragmented edaphic environments, where the latter observation was
probably due to limited dispersal/colonization opportunities or radiations of taxa in
allopatry. Pollination seems of importance in determining species rarity: sexual
deception had a significantly higher incidence of rarity than food-rewarding taxa. The
high incidence of rarity in sexually deceptive taxa could be due to either low fruit set
or the risk of specializing on a single pollinator species. They also concluded that
mycorrhizal infection could not explain the species rarity. It was clearly
demonstrated the role of pollinator species. Also this study applied GIS approach for
spatial analyses, and well demonstrated the spatial patterns of species commonness
and rarity. One may question their conclusion that fungal infection was not
important, due to a lack of fine-scaled measurement of fungal infection.
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[11] Kelly, C.K. et al. (2001). Investigations in commonness and rarity: a comparative
analysis of co-occurring, congeneric Mexican trees. Ecol. Lett. 4: 618-627.
This study focus on comparison for congeneic pairs in 12 tree species in Mexico.
They measured the population size distribution and tested the demographic changes
among common and rare species. They found common species showed more smooth
distribution in population size (low deviation of observed size from expected), with
contrast rare species experienced more fluctuation in their size distribution. Their
evidence pointed to the effects of recruitment upon patterns of being common and
rare. Although the focus is to looked demographic structures, it would be helpful to
present available information on their morphological traits if possible, for instance
how similar in their morphology and reproductive traits. Also direct measurements
on recruitment will be needed to draw an firm conclusion on effects of recruitment.
[12] Münzbergova, Z. (2005) Determinants of species rarity: population growth rates
of species sharing the same habitat. Am. J. Bot., 92, 1987–1994.
Although species traits are believed to be responsible to their occurrence and
commonness, demographic process is another key aspect to approach the question
why species remain common and rare. This study is one of examples compare
congeneric common versus rare species for mechanistic explanation. Life cycles of
one rare and one common Cirsium species sharing the same habitat were compared.
A systematic survey covered their distribution ranges for 13 C. pannonicum
populations (declined recently), and more than 90 C. acaule populations, and based
on Rabinowitz’s classification of rarity, both species would fall into the category of
wide ranges, large populations, and relatively narrow habitats. The distribution range
as well as range of habitat types are, however, somewhat narrower in the rare species.
The entire characteristics of life cycle were measured, including population growth
rate, seeds production and germination, and seed predation etc. Population growth
rate was slightly lower in the rare species, translating into a large difference in local
extinction probability. Seed predation intensity did not differ between species.
However, with prediction, common species showed less negative effects on seed
production and growth rate, therefore seed predation is the key factor causing a lower
population growth rate in the rare species. The study provided evidence on factors of
population growth rate (not ecological traits) that were responsible for commonness
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or rarity. They compared the demographic characteristics for congenic species based
on complete life cycle, not single traits, thus the gave unequivocal evidence of interspecies difference that ultimately manifested to their local abundance. Also detailed
information on demographic parameters is particular informative to direct
conservation decision, but on the other hand fine-scaled information will be at cost of
spending more time to collect field data.
[13] Cole, C.T. (2003) Genetic variation in rare and common plants. Ann. Rev. Ecol.
Evol. Syst., 34, 213–237
A meta-analyses on genetic variation and differentiation between common and rare
congeneric plant species were conducted, and summarizes the results from 144 reports
for 95 rare taxa and 152 corresponding common taxa. The purpose is to investigate
effects of small population size upon genetic structures. The isozyme variation in
these 247 plant species is summarized as 57 generic-level comparisons of rare and
common species. All species-level measures of variation and mean population-level
measures show reduction in rare species, but FIS and FST did not differ significantly,
reflecting the similarity of breeding system in congeneric species. The reduction in
gene flow among populations of rare species was significant. While not all measures
of genetic variation show reductions in all rare species, this report demonstrates that
the lower genetic variations in rare species. Genetic diversity and variability is
important aspects to study rare species, and this study highlight the possibility that
rare species were more impaired by low genetic diversity due to small population size.
Given the information provided, it is yet not clear whether low level of genetic
diversity is responsible to restricted distribution or low abundance of rare species,
apparently more studies that are explicitly designed to test this hypothesis is needed.
[14] Honnay, O. & Jacquemyn, H. (2007). Susceptibility of common and rare plant
species to the genetic consequences of habitat fragmentation. Conserv. Biol. 21: 823–
83.
The specific research questions are 1) whether small, fragmented plant populations
are more genetically impoverished compared with larger populations, and 2) whether
rare species are more vulnerable to habitat-fragmentation-mediated loss of genetic
diversity, how mating systems and longevity affect the relationships between
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population size and genetic variability. They addressed these questions with metaanalyses covering 53 publication and 52 species. Alleles lost through habitat
fragmentation and population size reduction were mainly those initially present at low
densities, and genetic bottle neck and drift play important role in genetic
impoverishment in plants. Genetic erosion is similarly likely in common and rare
species in response to habitat fragmentation. Genetic variability in self-compatible
species are less vulnerable to habitat fragmentation. The most important result is that
the common species are as vulnerable as rare species when affected by genetic
erosion associated with habitat fragmentation.
[15] Lososová, Z., Chytrý, M. & Kühn, I. (2008). Plant attributes determining the
regional abundance of weeds on central European Arable land. J. Biogeogr, 35, 177–
187.
The occurrence frequencies (regional abundance) of 381 weed species were examined
across Czech Republic, and correlated with their ecological traits, functional traits,
geographic range, and habitat use with regression tree. Associations of regional
abundance to phylogenetic relatedness and traits are compared. Rarity showed higher
correlation to ecological taints than phylogenetic relatedness, and phylogenetic
relatedness did not result in apparent changes on the association between rarity and
ecological traits, leading to them to conclude that the biological traits that are
responsible to cool weather adaptation is of particular importance in species
abundance in this study system. This study is the only example I managed to find that
directly compared three components environmental variables (estimated fundamental
niche), species traits, as well as phylogenetics. They also suggest their relative
importance, and which traits are of more interests varied among ecosystems and
organism groups.
In summary, like many biological and biogeographic topics regarding species relative
abundance (e.g. assembly-rules versus neutral theory), mechanistic understanding
beyond pattern description on commonness and rarity is still quite remote. One would
expect more research efforts toward mechanises understanding using various
disciplines. In particular, studies on aquatic systems, and studies involving population
structures or experimental approaches are extremely scarce. Regarding the patterns
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commonness and rarity showed significant variations between low versus high
diversity regions, as well as aquatic and terrestrial species, however large-scaled
studies conducted for aquatic taxa are too few to make informative generalization.
Hypothesis-testing frameworks that designed to explore cause-and-effect on species
commonness and rarity are desirable, either from spatial analyses or field
experiments. Phylogeny, demographic structures and population genetics has shed a
light on species rarity, but not many studies have been conducted applying those
methodologies. It is also interesting to compare population structures, trait values and
environmental variables among congeneric common versus rare species along their
distribution range.