The butterfly genus Heliconius contains some species that are extremely difficult to tell apart.
In biology, a species complex is a group of closely related
organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The
taxa in the complex may be able to hybridize readily with each other, further blurring any distinctions. Terms that are sometimes used synonymously but have more precise meanings are cryptic species for two or more species hidden under one species name, sibling species for two (or more) species that are each other's closest relative, and species flock for a group of closely related species that live in the same habitat. As informal
taxonomic ranks, species group, species aggregate, macrospecies, and superspecies are also in use.
Two or more taxa that were once considered
conspecific (of the same species) may later be subdivided into
infraspecific taxa (taxa within a species, such as bacterial
strains or plant
varieties), which may be a complex ranking but it is not a species complex. In most cases, a species complex is a
monophyletic group of species with a common ancestor, but there are exceptions. It may represent an early stage after
speciation in which the species were separated for a long time period without evolving
Hybrid speciation can be a component in the evolution of a species complex.
A species complex is typically considered as a group of close, but distinct species. Obviously, the concept is closely tied to the definition of a species. Modern biology understands a species as "separately evolving
metapopulationlineage" but acknowledges that the
criteria to delimit species may depend on the group studied. Thus, many traditionally defined species, based only on morphological similarity, have been found to be several distinct species when other criteria, such as genetic differentiation or
reproductive isolation, are applied.
A more restricted use applies the term to a group of species among which
hybridisation has occurred or is occurring, which leads to intermediate forms and blurred species boundaries. The informal classification, superspecies, can be exemplified by the
grizzled skipper butterfly, which is a superspecies that is further divided into three subspecies.
The Rubyspot damselfly
Hetaerina americana is suspected to be a cryptic complex with at least one other species of Rubyspot.
Several terms are used synonymously for a species complex, but some of them may also have slightly different or narrower meanings. In the
nomenclature codes of zoology and bacteriology, no
taxonomic ranks are defined at the level between
subgenus and species, but the botanical code defines four ranks below subgenus (section, subsection, series and subseries). Different informal taxonomic solutions have been used to indicate a species complex.
Also called physiologic race (uncommon). This describes "distinct species that are erroneously classified (and hidden) under one species name". More generally, the term is often applied when species, even if they are known to be distinct, cannot be reliably distinguished by morphology. The usage physiologic race is not to be confused with physiological race.
Also called aphanic species. This term, introduced by
Ernst Mayr in 1942, was initially used with the same meaning as cryptic species, but later authors emphasized the common phylogenetic origin. A recent article defines sibling species as "cryptic
sister species", "two species that are the closest relative of each other and have not been distinguished from one another taxonomically".
Also called species swarm. This refers to "a monophyletic group of closely related species all living in the same ecosystem". Conversely, the term has also been applied very broadly to a group of closely related species than can be variable and widespread. Not to be confused with a
Mixed-species foraging flock, a behavior in which birds of different species feed together.
Sometimes used as an informal rank for a species complex around one "representative" species. Popularized by
Bernhard Rensch and later
Ernst Mayr, with the initial requirement that species forming a superspecies must have
allopatric distributions. For the component species of a superspecies, allospecies was proposed.
Used for a species complex, especially in plant taxa where
apomixis are common. Historical synonyms are species collectiva, introduced by
Adolf Engler, conspecies, and grex. Components of a species aggregate have been called segregates or microspecies. Used as abbreviation agg. after the
binomial species name.
Distinguishing close species within a complex requires the study of often very small differences. Morphological differences may be minute and visible only by the use of adapted methods, such as
microscopy. However, distinct species sometimes have no morphological differences. In those cases, other characters, such as in the species'
karyology, may be explored. For example,
territorial songs are indicative of species in the
treecreepers, a bird genus with few morphological differences. Mating tests are common in some groups such as fungi to confirm the
reproductive isolation of two species.
A species complex typically forms a
monophyletic group that has diversified rather recently, as is shown by the short branches between the species A–E (blue box) in this
Species forming a complex have typically diverged very recently from each other, which sometimes allows the retracing of the process of
speciation. Species with differentiated populations, such as
ring species, are sometimes seen as an example of early, ongoing speciation: a species complex in formation. Nevertheless, similar but distinct species have sometimes been isolated for a long time without evolving differences, a phenomenon known as "morphological stasis". For example, the Amazonian frog Pristimantis ockendeni is actually at least three different species that diverged over 5 million years ago.
Stabilizing selection has been invoked as a force maintaining similarity in species complexes, especially when they adapted to special environments (such as a host in the case of symbionts or extreme environments). This may constrain possible directions of evolution; in such cases, strongly divergent selection is not to be expected. Also, asexual reproduction, such as through
apomixis in plants, may separate lineages without producing a great degree of morphological differentiation.
Possible processes explaining similarity of species in a species complex: a – morphological stasis b –
Sources differ on whether or not members of a species group share a
range. A source from
Iowa State University Department of
Agronomy states that members of a species group usually have partially overlapping ranges but do not
interbreed with one another.A Dictionary of Zoology (
Oxford University Press 1999) describes a species group as complex of related species that exist
allopatrically and explains that the "grouping can often be supported by experimental crosses in which only certain pairs of species will produce
hybrids." The examples given below may support both uses of the term "species group."
Often, such complexes do not become evident until a new species is introduced into the system, which breaks down existing species barriers. An example is the introduction of the
Spanish slug in
Northern Europe, where interbreeding with the local
black slug and
red slug, which were traditionally considered clearly separate species that did not interbreed, shows that they may be actually just subspecies of the same species.
Where closely related species co-exist in
sympatry, it is often a particular challenge to understand how the similar species persist without outcompeting each other.
Niche partitioning is one mechanism invoked to explain that. Indeed, studies in some species complexes suggest that species divergence have gone in par with ecological differentiation, with species now preferring different microhabitats.
Similar methods also found that the Amazonian frog Eleutherodactylus ockendeni is actually at least three different species that diverged over 5 million years ago.
It has been suggested that cryptic species complexes are very common in the marine environment. That suggestion came before the detailed analysis of many systems using DNA sequence data but has been proven to be correct. The increased use of DNA sequence in the investigation of organismal diversity (also called
DNA barcoding) has led to the discovery of a great many cryptic species complexes in all habitats. In the marine bryozoan Celleporella hyalina, detailed morphological analyses and mating compatibility tests between the isolates identified by DNA sequence analysis were used to confirm that these groups consisted of more than 10 ecologically distinct species, which had been diverging for many millions of years.
Evidence from the identification of cryptic species has led some[who?] to conclude that current estimates of global species richness are too low.
Pests, species that cause diseases and their vectors, have direct importance for humans. When they are found to be cryptic species complexes, the ecology and the virulence of each of these species need to be re-evaluated to devise appropriate control strategies. Examples are cryptic species in the
malaria vector genus of mosquito, Anopheles, the fungi causing
cryptococcosis, and sister species of Bactrocera tryoni, or the Queensland fruit fly. That pest is indistinguishable from two sister species except that B. tryoni inflicts widespread, devastating damage to Australian fruit crops, but the sister species do not.
When a species is found to be several phylogenetically distinct species, each typically has smaller distribution ranges and population sizes than had been reckoned. The different species can also differ in their ecology, such as by having different breeding strategies or habitat requirements, which must be taken into account for appropriate management. For example,
giraffe populations and subspecies differ genetically to such an extent that they may be considered species. Although the giraffe, as a whole, is not considered to be threatened, if each cryptic species is considered separately, there is a much higher level of threat.
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