Cryptospore
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Cryptospores are microscopic
fossilized A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s produced by
embryophytes The embryophytes () are a clade of plants, also known as Embryophyta (Plantae ''sensu strictissimo'') () or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophy ...
(land plants). They first appear in the
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
record during the middle of the
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...
period, as the oldest fossil evidence for the colonization of land by plants. A similar (though broader) category is miospores, a term generally used for spores smaller than 200
μm The micrometre (Commonwealth English as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American English), also commonly known by the non-SI term micron, is a unit of length in the International System ...
. Both cryptospores and miospores are types of
palynomorphs Palynology is the study of microorganisms and microscopic fragments of mega-organisms that are composed of acid-resistant organic material and occur in sediments, sedimentary rocks, and even some metasedimentary rocks. Palynomorphs are the mic ...
. Cryptospores, which occur as permanent tetrads, dyads, or hilate monads, sometimes with additional wall envelopes, dominated fossil assemblages for approximately 60 million years, first appearing around 470 million years ago during the Ordovician Period. They underwent rapid diversification during what Jane Gray (1993) called the Eoembryophytic epoch, but experienced an abrupt decline in diversity and abundance around 410 Ma during the latest
Lochkovian The Lochkovian is one of three faunal stages in the Early Devonian Epoch. It lasted from 419.2 ± 3.2 million years ago to 410.8 ± 2.8 million years ago. It marked the beginning of the Devonian Period, and was followed by the Pragian Stage. It is ...
(Early Devonian), with only a few forms persisting into the Emsian period. In contrast, trilete monads began diversifying around 430 Ma in the latter Silurian Period and eventually became the dominant element in dispersed spore assemblages. While trilete monads are generally associated with vascular plants, cryptospores lack close modern analogues (except possibly in some liverworts), making the identification of their parent plants one of evolutionary botany's significant unresolved problems.


Evidence that cryptospores derive from land plants


Occurrence

Cryptospores are generally found in non-marine
rock Rock most often refers to: * Rock (geology), a naturally occurring solid aggregate of minerals or mineraloids * Rock music, a genre of popular music Rock or Rocks may also refer to: Places United Kingdom * Rock, Caerphilly, a location in Wale ...
s and decrease in abundance with distance offshore. This suggests that any cryptospores found in the marine environment were transported there by the wind from the land, rather than originating from the marine environment.


Wall ultrastructure

The walls of cryptospores consist of many lamellae (thin sheets).
Liverwort Liverworts are a group of non-vascular land plants forming the division Marchantiophyta (). They may also be referred to as hepatics. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry ...
s, thought to be the most primitive
land plants The embryophytes () are a clade of plants, also known as Embryophyta (Plantae ''sensu strictissimo'') () or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophy ...
, also have this spore wall
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
.


Chemical composition

(Some) cryptospores are composed of sporopollenin and have the same chemical makeup as co-occurring trilete spores.


Other information

Recently, fossils of plant
sporangia A sporangium (from Late Latin, ; : sporangia) is an enclosure in which spores are formed. It can be composed of a unicellular organism, single cell or can be multicellular organism, multicellular. Virtually all plants, fungus, fungi, and many ot ...
have been found in
Oman Oman, officially the Sultanate of Oman, is a country located on the southeastern coast of the Arabian Peninsula in West Asia and the Middle East. It shares land borders with Saudi Arabia, the United Arab Emirates, and Yemen. Oman’s coastline ...
with cryptospores showing concentric lamellae in their walls, similar to
liverworts Liverworts are a group of non-vascular plant, non-vascular embryophyte, land plants forming the division Marchantiophyta (). They may also be referred to as hepatics. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in wh ...
. The earliest known cryptospores are from Middle
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...
(
Dapingian The Dapingian is the third stage of the Ordovician system and the first stage of the Middle Ordovician series. It is preceded by the Floian and succeeded by the Darriwilian. The base of the Dapingian (and the top of the Floian) is defined as t ...
) strata of
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
. Spores from the Lindegård
Mudstone Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from ''shale'' by its lack of fissility.Blatt, H., and R.J. Tracy, 1996, ''Petrology.'' New York, New York, ...
(late
Katian The Katian is the second stage of the Upper Ordovician. It is preceded by the Sandbian and succeeded by the Hirnantian Stage. The Katian began million years ago and lasted for about 7.6 million years until the beginning of the Hirnantian mill ...
–early
Hirnantian The Hirnantian is the final internationally recognized stage of the Ordovician Period of the Paleozoic Era. It was of short duration, lasting about 2.1 million years, from to Ma (million years ago). The early part of the Hirnantian was char ...
) represent the earliest record of early
land plant The embryophytes () are a clade of plants, also known as Embryophyta (Plantae ''sensu strictissimo'') () or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophyt ...
spores In biology, a spore is a unit of sexual (in fungi) or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plant ...
from
Sweden Sweden, formally the Kingdom of Sweden, is a Nordic countries, Nordic country located on the Scandinavian Peninsula in Northern Europe. It borders Norway to the west and north, and Finland to the east. At , Sweden is the largest Nordic count ...
and possibly also from
Baltica Baltica is a paleocontinent that formed in the Paleoproterozoic and now constitutes northwestern Eurasia, or Europe north of the Trans-European Suture Zone and west of the Ural Mountains. The thick core of Baltica, the East European Craton, i ...
and implies that
land plants The embryophytes () are a clade of plants, also known as Embryophyta (Plantae ''sensu strictissimo'') () or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophy ...
had migrated to the
palaeocontinent A paleocontinent or palaeocontinent is a distinct area of continental crust that existed as a major landmass in the geological past. There have been many different landmasses throughout Earth's time. They range in sizes; some are just a collection ...
Baltica Baltica is a paleocontinent that formed in the Paleoproterozoic and now constitutes northwestern Eurasia, or Europe north of the Trans-European Suture Zone and west of the Ural Mountains. The thick core of Baltica, the East European Craton, i ...
by at least the Late
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...
. This discovery reinforces the earlier suggestion that the migration of land plants from northern
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
to
Baltica Baltica is a paleocontinent that formed in the Paleoproterozoic and now constitutes northwestern Eurasia, or Europe north of the Trans-European Suture Zone and west of the Ural Mountains. The thick core of Baltica, the East European Craton, i ...
in the Late
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...
was facilitated by the northward migration of
Avalonia Avalonia was a microcontinent in the Paleozoic era. Crustal fragments of this former microcontinent are terranes in parts of the eastern coast of North America: Atlantic Canada, and parts of the East Coast of the United States, East Coast of the ...
, which is evidenced by the co-occurrence of reworked, Early–Middle
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...
acritarchs Acritarchs () are organic microfossils, known from the Precambrian to the present. The classification is a catch all term used to refer to any organic microfossils that cannot be assigned to other groups. Their diversity reflects major ecologi ...
, possibly suggesting an Avalonian provenance in a
foreland basin A foreland basin is a structural basin that develops adjacent and parallel to a mountain belt. Foreland basins form because the immense mass created by crustal thickening associated with the evolution of a mountain belt causes the lithospher ...
system.


See also

*
Paleobotany Paleobotany or palaeobotany, also known as paleophytology, is the branch of botany dealing with the recovery and identification of plant fossils from geological contexts, and their use for the biological reconstruction of past environments ( pal ...
*
Palynology Palynology is the study of microorganisms and microscopic fragments of mega-organisms that are composed of acid-resistant organic material and occur in sediments, sedimentary rocks, and even some metasedimentary rocks. Palynomorphs are the mic ...


References


Further reading

* {{cite journal , last1=Edwards , first1=D. , last2=Morris , first2=J. L. , last3=Richardson , first3=J. B. , last4=Kenrick , first4=P. , year=2014 , title=Cryptospores and cryptophytes reveal hidden diversity in early land floras , journal=New Phytologist , volume=202 , number=1 , pages=50–78 , doi=10.1111/nph.12645, pmid=24410730 , url=https://orca.cardiff.ac.uk/57757/1/EdwardsD.EARTH3.pdf Fossil record of plants Microfossils Ordovician plants Paleobotany Paleozoic life Palynology Silurian life