Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Polycarpon tetraphyllum
(fourleaf allseed)

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Datasheet

Polycarpon tetraphyllum (fourleaf allseed)

Summary

  • Last modified
  • 19 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Polycarpon tetraphyllum
  • Preferred Common Name
  • fourleaf allseed
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • P. tetraphyllum is a small and relatively insignificant annual plant. It is a widespread weed of cultivated and waste areas. Thought to be native originally to the Mediterranean region, P. tetratphyllu...

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Pictures

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PictureTitleCaptionCopyright
Polycarpon tetraphyllum (fourleaf manyseed); vegetative plant showing habit.
TitleHabit
CaptionPolycarpon tetraphyllum (fourleaf manyseed); vegetative plant showing habit.
Copyright©Trevor James/Hamilton, New Zealand-2014
Polycarpon tetraphyllum (fourleaf manyseed); vegetative plant showing habit.
HabitPolycarpon tetraphyllum (fourleaf manyseed); vegetative plant showing habit.©Trevor James/Hamilton, New Zealand-2014
Polycarpon tetraphyllum (fourleaf manyseed); habit, whole plant flowering and fruiting.
TitleHabit
CaptionPolycarpon tetraphyllum (fourleaf manyseed); habit, whole plant flowering and fruiting.
Copyright©Trevor James/Hamilton, New Zealand-2014
Polycarpon tetraphyllum (fourleaf manyseed); habit, whole plant flowering and fruiting.
HabitPolycarpon tetraphyllum (fourleaf manyseed); habit, whole plant flowering and fruiting.©Trevor James/Hamilton, New Zealand-2014
Polycarpon tetraphyllum (fourleaf manyseed); massed inflorescences.
TitleInflorescences
CaptionPolycarpon tetraphyllum (fourleaf manyseed); massed inflorescences.
Copyright©Trevor James/Hamilton, New Zealand-2014
Polycarpon tetraphyllum (fourleaf manyseed); massed inflorescences.
InflorescencesPolycarpon tetraphyllum (fourleaf manyseed); massed inflorescences.©Trevor James/Hamilton, New Zealand-2014
Polycarpon tetraphyllum (fourleaf manyseed); close-up of open and closed flowers.
TitleFlowers
CaptionPolycarpon tetraphyllum (fourleaf manyseed); close-up of open and closed flowers.
Copyright©Trevor James/Hamilton, New Zealand-2014
Polycarpon tetraphyllum (fourleaf manyseed); close-up of open and closed flowers.
FlowersPolycarpon tetraphyllum (fourleaf manyseed); close-up of open and closed flowers.©Trevor James/Hamilton, New Zealand-2014

Identity

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Preferred Scientific Name

  • Polycarpon tetraphyllum (L.) L.

Preferred Common Name

  • fourleaf allseed

Other Scientific Names

  • Alsine polycarpa Crantz
  • Mollugo tetraphylla L.
  • Polycarpaea tetrphylla (L.) E.H.L.Krause
  • Polycarpon tetraphyllum subsp. alsinifolium (Biv.) Ball
  • Polycarpon tetraphyllum subsp. diphyllum (Cav.) O.Bolos & Font Quer

International Common Names

  • English: four-leaf allseed; four-leaved allseed

Local Common Names

  • : allseed; fourleaf manyseed; polycarpon
  • : polycarpe a 4 feuilles
  • Germany: vierblattriges Nagelkraut
  • Italy: migliarina a 4 foglie
  • Netherlands: kransmuur
  • South Africa: Naaldvrug
  • Sweden: Kransort; tusenfro

Summary of Invasiveness

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P. tetraphyllum is a small and relatively insignificant annual plant. It is a widespread weed of cultivated and waste areas. Thought to be native originally to the Mediterranean region, P. tetratphyllum is now present on every continent except Antarctica. It is regarded as an invasive species in Australia, New Zealand, Japan, parts of South America and the USA, including Hawaii.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Caryophyllales
  •                         Family: Caryophyllaceae
  •                             Genus: Polycarpon
  •                                 Species: Polycarpon tetraphyllum

Notes on Taxonomy and Nomenclature

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P. tetraphyllum is described as one of about 16 species of the genus Polycarpon (Mabberley, 1997); however, Kool et al. (2007) pointed out the extensive synonymy, especially in the Polycarpon tetraphyllum group, suggesting that the number of taxa in the genus should be as few as eight. The Polycarpon taxa distributed in the Mediterranean region are largely ecotypes adapted to different environmental conditions rather than separate species (Kool et al., 2007). The same authors suggested that P. depressum, which grows in a relatively limited area in California and Mexico, may be the result of an earlier dispersal of a P. tetraphyllum lineage which had been isolated for some time and become adapted to the local environment. 

Description

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Modified from Webb et al. (1988):

Annual, rarely biennial, or possibly perennial, taprooted herb. Stems glabrous, geniculate, erect to ascending, much-branched, 5-20 cm long. Leaves green or reddish, rarely purplish, glabrous, in whorls of 4 or rarely in opposite pairs, obovate to suborbicular, obtuse or shortly apiculate, petiolate, sometimes falling at fruiting, (2)-5-10-(15) × (1)-3-8-(10) mm. Stipules scarious, triangular to ovate, acuminate, 2-3 × 1-1.5 mm. Inflorescence many-flowered, lax or rarely compact. Pedicels about equal to calyx, equal or greater than the ovate acuminate scarious bracts. Sepals apiculate, hooded, green or reddish, rarely purplish, with scarious margins, 2-2.5 × 1 mm. Petals white, narrow-elliptic, entire, 1 mm long. Stamens 3-5, 0.5-0.7 mm long. Ovary 0.5 mm long; style 0.3-0.4 mm long, 3-lobed, deciduous. Capsule ovoid, about 2 mm long. Seeds curved, brown, finely papillate.

Plant Type

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Biennial
Broadleaved
Herbaceous
Seed propagated

Distribution

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The exact native range of P. tetraphyllum is obscure, but most authorities suggest it is native to southern Europe and the Mediterranean region (Weeds of Australia, 2013). Using DNA sequence data, Kool et al. (2007) found that the P. tetraphyllum clade is most diverse in the Mediterranean region, where they deduced that it probably originated. Weedy forms of the species have been spread to every continent except Antarctica.

P. tetraphyllum has been introduced to India, Japan, the Americas, Australia and New Zealand, parts of Europe and a few countries in sub-Saharan Africa.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

AzerbaijanPresentNativeEuro+Med Plantbase, 2013
IndiaPresentIntroducedBiodiversity India, 2013Growing as a weed in tea plantations and roadsides in the High Ranges
-KeralaPresentIntroducedBiodiversity India, 2013
-Tamil NaduPresentIntroducedBiodiversity India, 2013
IranPresentNativeUSDA-ARS, 2013
IsraelPresentNativeUSDA-ARS, 2013
JapanPresentIntroduced Invasive Mito and Uesugi, 2004
JordanPresentNativeEuro+Med Plantbase, 2013
KuwaitPresentNativeUSDA-ARS, 2013
LebanonPresentNativeEuro+Med Plantbase, 2013
OmanPresentNativeUSDA-ARS, 2013
QatarPresentNativeUSDA-ARS, 2013
Saudi ArabiaPresentNativeEuro+Med Plantbase, 2013
SyriaPresentNativeEuro+Med Plantbase, 2013
TurkeyPresentNativeEuro+Med Plantbase, 2013
YemenPresentNativeUSDA-ARS, 2013

Africa

AlgeriaPresentNativeEuro+Med Plantbase, 2013
Cape VerdePresentDiniz, 1995Likely to be native
EgyptPresentNativeEuro+Med Plantbase, 2013
EritreaPresentNativeUSDA-ARS, 2013
EthiopiaPresentIntroducedUSDA-ARS, 2013
LibyaPresentNativeEuro+Med Plantbase, 2013
MoroccoPresentNativeEuro+Med Plantbase, 2013
South AfricaPresentIntroduced Invasive Randall, 2012
Spain
-Canary IslandsPresentNativeEuro+Med Plantbase, 2013
TunisiaPresentNativeEuro+Med Plantbase, 2013

North America

CanadaPresentIntroducedGBIF, 2013
-British ColumbiaPresentIntroducedGBIF, 2013
MexicoPresentIntroducedMissouri Botanical Garden, 2013
USAPresent, few occurrencesIntroducedNew England Wild Flower Society, 2013Mainly in the south and west
-AlabamaPresentIntroducedUSDA-ARS, 2013
-CaliforniaPresentIntroducedGBIF, 2013
-FloridaPresentIntroducedGBIF, 2013
-GeorgiaPresentIntroducedUSDA-ARS, 2013
-HawaiiPresent, few occurrencesIntroduced Invasive PIER, 2013
-MassachusettsPresent, few occurrencesIntroducedNew England Wild Flower Society, 2013Only collected in Massachusetts in association with the waste dump of the 19th century wool-carding factory
-North CarolinaPresentIntroducedGBIF, 2013
-OregonPresentIntroducedGBIF, 2013
-PennsylvaniaPresentIntroducedGBIF, 2013
-South CarolinaPresentIntroducedUSDA-NRCS, 2013
-TexasPresentIntroducedGBIF, 2013

Central America and Caribbean

Costa RicaPresentIntroducedUSDA-ARS, 2013
HaitiPresentIntroducedMissouri Botanical Garden, 2013

South America

ArgentinaPresentIntroducedMissouri Botanical Garden, 2013
BrazilPresentIntroducedReflora, 2013
-ParaPresentIntroducedReflora, 2013
-ParanaPresentIntroducedReflora, 2013
-Rio de JaneiroPresentIntroducedReflora, 2013
-Rio Grande do SulPresentIntroducedReflora, 2013
-Santa CatarinaPresentIntroducedReflora, 2013
-Sao PauloPresentIntroducedReflora, 2013
ChilePresentIntroduced Invasive PIER, 2013
-Easter IslandPresentIntroducedPIER, 2013
EcuadorPresentIntroduced Invasive PIER, 2013
PeruPresentIntroducedPIER, 2013
UruguayPresentIntroducedRios et al., 2008; Randall, 2012
VenezuelaPresentIntroducedMissouri Botanical Garden, 2013

Europe

AlbaniaPresentNativeUSDA-ARS, 2013
BelgiumPresentIntroducedDAISIE, 2013
BulgariaPresentNativeEuro+Med Plantbase, 2013
CroatiaPresentNativeEuro+Med Plantbase, 2013
CyprusPresentNativeEuro+Med Plantbase, 2013
DenmarkPresentIntroducedDAISIE, 2013
FinlandPresentIntroducedKurtto and Lahti, 1987
FrancePresent, few occurrencesNativeHYPPA, 2013Esp. vineyards, orchards, crops in south
GermanyPresentNativeEuro+Med Plantbase, 2013
ItalyPresentNativeEuro+Med Plantbase, 2013
MacedoniaPresentNativeEuro+Med Plantbase, 2013
MaltaPresentNativeEuro+Med Plantbase, 2013
MontenegroPresentNativeUSDA-ARS, 2013
NetherlandsPresentIntroducedDAISIE, 2013
PortugalPresentIntroducedEuro+Med Plantbase, 2013
-AzoresPresentIntroducedEuro+Med Plantbase, 2013
-MadeiraPresentNativeEuro+Med Plantbase, 2013
RomaniaPresentNativeEuro+Med Plantbase, 2013
SlovakiaPresentIntroducedEuro+Med Plantbase, 2013
SloveniaPresentNativeEuro+Med Plantbase, 2013
SpainPresentNative Invasive HYPPA, 2013Esp. in olives and vineyards, also in cereals & sugarbeet
-Balearic IslandsPresentNativeEuro+Med Plantbase, 2013
SwedenPresentIntroducedDAISIE, 2013
SwitzerlandPresentNativeEuro+Med Plantbase, 2013
UKPresent, few occurrencesNativeStace, 2010
Yugoslavia (former)PresentNativeEuro+Med Plantbase, 2013

Oceania

AustraliaWidespreadIntroduced Invasive Weeds of Australia, 2013
-Australian Northern TerritoryWidespreadIntroduced Invasive Weeds of Australia, 2013Southern parts
-New South WalesWidespreadIntroduced Invasive Weeds of Australia, 2013
-QueenslandWidespreadIntroduced Invasive Weeds of Australia, 2013Southern and central areas
-South AustraliaWidespreadIntroduced Invasive Weeds of Australia, 2013
-TasmaniaWidespreadIntroduced Invasive Weeds of Australia, 2013
-VictoriaWidespreadIntroduced Invasive Weeds of Australia, 2013
-Western AustraliaPresentIntroduced Invasive Weeds of Australia, 2013Southern and western areas
New CaledoniaPresentIntroducedPIER, 2013
New ZealandWidespreadIntroduced Invasive Webb et al., 1988
Norfolk IslandPresentIntroducedWeeds of Australia, 2013

History of Introduction and Spread

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According to Thieret and Rabeler (2013), the spread of P. tetraphyllum to North America was partly due to its storage in ships’ ballast. Its seed is extremely small and could easily have been transported to North America and elsewhere as a contaminant in soil, dirt or vegetable matter taken there deliberately or accidently.

Historic records of P. tetraphyllum are known from Massachusetts in 1917 and Pennsylvania in the 1890s (Thieret and Rabeler, 2013).

According to Australia’s Virtual Herbarium (2013) the species was first found in Victoria in 1853 and in Tasmania in 1854. In New Zealand it was first recorded in 1854 (Webb et al., 1998).

In the UK, P. tetraphyllum has been known in Dorset since 1770 and south Devon since 1778 (Biological Records Centre, 2013). Populations can fluctuate dramatically and P. tetraphyllum has been rediscovered at some sites after an apparent absence of many years.

P. tetraphyllum was introduced to Belgium as a weed in containers with Mediterranean plants, and is now expanding rapidly in warmer microclimates of urban areas in that country.

Risk of Introduction

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P. tetraphyllum has very small, dust-like seeds produced in large numbers. The transport of seeds to other countries or other parts of countries where P. tetraphyllum does not yet occur is therefore likely. Seeds can be transported as a contaminant in agricultural or horticultural seeds, in other agricultural produce or among the dust that tends to travel with passengers, their goods and luggage.

Habitat

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P. tetraphyllum is generally a small weed of waste places, cultivated land and open pastures. In its native Mediterranean, P. tetraphyllum is found in woodlands and shrublands, deserts and montane vegetation (Danin, 2013).

In California, Hartman and Rabeler (2012) described its habitat as ‘disturbed shaded areas [and] roadsides’ below 450 m.

Biological Records Centre (2013) described its habitat in the UK as ‘open, sunny sites that are droughted in summer and relatively frost-free in winter. It grows with other therophytes on steep south-facing banks, on compacted shingle or sand, in bulb-fields and gardens and at the base of roadside walls.’ Clapham et al. (1962) described its habitat in the UK as sandy and waste places in the southeast (Cornwall, south Devon and Dorset), but added that suitable habitats for P. tetraphyllum in the UK were rare.

In New Zealand it is found growing in gravelly or sandy sites, coastal and inland areas, riverbeds, roadsides, railway ballast, seashore, waste land, cultivated land and grassland (Webb et al., 1988).

The ability of annual plants, including P. tetraphyllum, to colonise open habitats (because of their annual nature and wind-dispersed seed) and to capture resources effectively (because of the flat architecture of their rosettes) means they are more likely to occur in disturbed sites in temperate grasslands of the New England Tablelands of Australia (McIntyre et al., 1995). Disturbance by heavy grazing can increase the proportion of such annuals in these grasslands.

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial ‑ Natural / Semi-naturalHigh altitudes, uplands Principal habitat
 
Terrestrial – ManagedCultivated / agricultural land Present, no further details
Managed grasslands (grazing systems) Principal habitat
Rail / roadsides Present, no further details
Natural forests Principal habitat
Natural grasslands Present, no further details
Riverbanks Present, no further details
Scrub / shrublands Principal habitat
Deserts Principal habitat
Arid regions Principal habitat
Littoral
Coastal areas Present, no further details

Biology and Ecology

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Genetics

2n=32, 48, or 64 (Hartman and Rabeler, 2012).

Reproductive biology

The flowers of P. tetraphyllum are homogynous and automatically self-pollinated, and often cleistogamous (Clapham et al., 1962). In the Mediterranean region, Danin (2013) reported it flowering in almost every month of the year. In Britain it flowers in June and July (Clapham et al., 1962). In Australia P. tetraphyllum flowers in spring and continues into summer (Herbiguide, 2013).

In Australia, seed germinates in autumn to winter and plants then grow over winter through to summer (Herbiguide, 2013).

Kew Royal Botanic Gardens (2013) quoted an average 1000 seed weight of P. tetraphyllum as 0.05g. The seeds showed 100% germination at 10o or 11oC in 8 or 12 hours day length, although their age when tested was not mentioned.

A seed bank in Spain dominated by tiny seeds of P. tetraphyllum and Vulpia spp. contained up to 126,600 seeds m-2 (Marañón, 1998; Olea and San Miguel-Ayanz, 2006). Lunt (1997) found seeds of P. tetraphyllum in the soil under intact remnants of Eucalyptus tereticornis woodland in temperate south east Australia, both in grassy forest or small grassland patches in the forest. P. tetraphyllum was not represented in the vegetation. More seeds were found in soil from the forest remnants than in soil from the grassland patches. Buisson et al (2006), working on abandoned fields in southern France, found seeds of P. tetraphyllum in few of the surface soil samples in winter (persistent seedbank), and even fewer in summer surface soil samples (transient seedbank).

Physiology and phenology

In Australia, P. tetraphyllum plants can be annual or biennial, with seed germinating in autumn to winter and the plants then growing over winter through to summer. They start flowering in spring and this continues into summer (Herbiguide, 2013).

Using old herbarium records, Buswell et al. (2011) investigated whether short-lived plants (including P. tetraphyllum) introduced to Australia before 1920 were evolving rapidly to the different conditions there. The only significant change in P. tetraphyllum was a reduction in plant height in specimens from the semi-arid region of New South Wales, but plant height in higher rainfall coastal New South Wales either did not change or increased over time. The authors suggested that this pattern ‘is consistent with the idea that the more arid environmental conditions in western New South Wales have selected for decreased height’ in this and a few other species.

Longevity

P. tetraphyllum is usually an annual, although there is a report of it as a short-lived perennial (Herbiguide, 2013). No information seems to be available on the longevity of seed in the soil.

Population size and structure

In Australia, P. tetraphyllum can produce a dense cover in moist, fertile areas (Herbiguide, 2013). Since individual plants tend to be quite small, this implies that plant densities can be very high in appropriate conditions. Gremmen and Halbertsma (2009) also found the species had high cover value at times in Tristan da Cunha, an island in the South Atlantic.

Means of Movement and Dispersal

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Natural dispersal (non-biotic)

The seeds of P. tetraphyllum are very small and light and can be carried by air currents. However, since the parent plant is small (under 15 cm tall), long-distance dispersal of its seeds may be limited.

Vector transmission (biotic)

Calviño-Cancela et al. (2006) found seedlings of P. tetraphyllum emerging from the droppings of emu (Dromaius novaehollandiae). Twigg et al. (2008) reported the occurrence of whole or identifiable fragments of seeds of Polycarpon (perhaps P. tetraphyllum) in rabbit (Oryctolagus cuniculus) faecal pellets.

Buisson et al. (2006) assessed above-ground vegetation, seed rain, ant-borne seeds and seed banks on three abandoned fields in southeast France, at the margins between these fields and a remnant patch of native steppe vegetation of high environment value. P. tetraphyllum was present in 95% of the 990 quadrats used, but its seeds were only found in 2.8 % of the 1287 seed traps and in 0.4 % of the ant pitfall traps. Ants therefore seem to play only a small part in seed dispersal.

Accidental introduction

The seeds of P. tetraphyllum are tiny and can easily contaminate agricultural or horticultural seed, produce, machinery, or amongst the dust that tends to travel with passengers, their goods and luggage, and could easily be taken to new countries or to different parts of countries where the plant already occurs.

P. tetraphyllum is a wool alien (carried in wool used in manufacturing processes); in Massachusetts, USA, it has only been collected ‘in association with the waste dump of a nineteenth-century wool-carding factory’ (GoBotany, 2013). Hoste and Verloove (2009) described it as a container alien; it was introduced to Belgium as a weed in containers with Mediterranean plants.

Intentional introduction

Intentional introduction is very unlikely as P. tetraphyllum seems to have neither medicinal nor herbal uses and is a very small and inconspicuous plant.

Environmental Impact

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Although usually considered a very small and insignificant weed, P. tetraphyllum can sometimes grow in very dense patches (Gremmen and Halbertsma, 2009; Herbiguide, 2013).

On the tiny South Atlantic island of Tristan da Cunha, Gremmen and Halbertsma (2009) reported that it is 'one of the species invading native dynamic habitats, thus contributing to the transformation of these plant communities into aliens-dominated vegetation.'

Social Impact

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P. tetraphyllum is often regarded as a minor weed in gardens and occasionally in lawns, where it may reduce their aesthetic value.

On the tiny south Atlantic island of Tristan da Cunha, Gremmen and Halbertsma (2009) reported that ‘in gardens and arable land, as well as along road verges, it [P. tetraphyllum] locally reaches very high cover values. It is an abundant weed in the [potato] patches, and has a clear negative impact on food production.’ 

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Pioneering in disturbed areas
  • Fast growing
  • Has high reproductive potential
  • Has propagules that can remain viable for more than one year
  • Has high genetic variability
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult to identify/detect as a commodity contaminant

Similarities to Other Species/Conditions

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In Australia, P. tetraphyllum resembles Crassula spp., Stellaria media, Cerastium spp., Portulaca spp., Anagallis arvenis, Montia spp., Sagina apetala and Euphorbia peplus (Herbiguide, 2013). All are low-growing, small weedy herbs, but most become taller or have much longer sprawling stems compared to P. tetraphyllum. Popay et al. (2010) identified Sagina apetala as the species most likely to be confused with P. tetraphyllum, but S. apetala has very narrow leaves in pairs and the flowers are less densely clustered than P. tetraphyllum.  

P. tetraphyllum var. diphyllum can be distinguished by leaves only paired (as opposed to being in fours), contracted inflorescences and 1-3 stamens, instead of 3-5 (Stace, 2010). 

Prevention and Control

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Physical/mechanical control

Where P. tetraphyllum is established, Herbiguide (2013) suggested preventing seed set to control the plant.

Biological control

P. tetraphyllum is generally considered too small and insignificant to merit any major biological control effort.

Chemical control

The Novachem Manual (2012) lists 2,4-D ester for use in pastures, acetochlor in maize crops or terbacil in some fruit crops. There seems to be little information elsewhere on chemical control, other than the ubiquitous use of glyphosate in total weed control.

Control by utilization

Herbiguide (2013) suggests grazing the area where P. tetraphyllum is established. However, heavy grazing can increase the proportion of P. tetraphyllum and other small annual species in grassland (McIntyre et al., 1995). 

Gaps in Knowledge/Research Needs

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Little information is available on the germination or longevity of seeds of P. tetraphyllum, nor on the slightly puzzling references to the plants being sometimes ‘perennial’ (Herbiguide, 2013). Nor is there information on how soon after germination the plants can flower and produce viable seed.

References

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Biodiversity India, 2013. India Biodiversity Portal. http://indiabiodiversity.org/

Biological Records Centre, 2013. Online Atlas of the British and Irish flora. Wallingford, UK: Biological Records Centre. http://www.brc.ac.uk/plantatlas/

Buisson E; Dutoit T; Torre F; Römermann C; Poschlod P, 2006. The implications of seed rain and seed bank patterns for plant succession at the edges of abandoned fields in Mediterranean landscapes. Agriculture, Ecosystems & Environment, 115(1/4):6-14.

Buswell JM; Moles AT; Hartley S, 2011. Is rapid evolution common in introduced plant species? Journal of Ecology (Oxford), 99(1):214-224. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1365-2745

Calviño-Cancela M; Dunn RR; Etten EJBvan; Lamont BB, 2006. Emus as non-standard seed dispersers and their potential for long-distance dispersal. Ecography, 29(4):632-640. http://www3.interscience.wiley.com/journal/118729914/abstract

Clapham AR; Tutin TG; Warburg EF, 1962. Flora of the British Isles. Second edition. Cambridge, UK: Cambridge University Press.

Cruz A; Botelho M; Silvestre J; Castro Rde, 2012. Soil management: introduction of tillage in a vineyard with a long-term natural cover. Ciência e Técnica Vitivinícola, 27(1):27-38.

DAISIE, 2013. Delivering Alien Invasive Species Inventories for Europe. DAISIE (online). www.europe-aliens.org

Danin A, 2013. Flora of Israel online. Jerusalem, Israel: The Hebrew University of Jerusalem. http://flora.huji.ac.il/browse.asp

Diniz MA, 1995. Cape Verde flora. Vascular plants. Caryophyllaceae. (Flora de Cabo Verde. Plantas vasculares. Caryophyllaceae.) Flora de Cabo Verde, No. 19. Lisboa, Portugal: Instituto de Investigação Científica Tropical, 26 pp.

Euro+Med Plantbase, 2013. The information resource for Euro-Mediterranean plant diversity. The information resource for Euro-Mediterranean plant diversity. unpaginated. http://ww2.bgbm.org/EuroPlusMed/

GBIF, 2013. Global Biodiversity Information Facility. Global Biodiversity Information Facility (GBIF). http://data.gbif.org/species/

Gibbs Russell GE; Welman WG; Reitief E; Immelman KL; Germishuizen G; Pienaar BJ; Wyk Mvan; Nicholas A, 1987. List of species of southern African plants. Memoirs of the Botanical Survey of South Africa, 2(1 & 2):1-152 & 1-270.

Gremmen N; Halbertsma RL, 2009. Alien plants and their impact on Tristan da Cunha. Part 2: Species accounts. Alien plants and their impact on Tristan da Cunha. http://herbaria.plants.ox.ac.uk/bol/Content/Groups/tristan/Resources/Tristan_Alien_Plant_Report_Part2.pdf

Hartman RL; Rabeler RK, 2012. Jepson Flora Project (eds.) Jepson eFlora. Jepson Herbarium. California, USA: University of California, Berkeley. http://ucjeps.berkeley.edu/jeps/

Herbiguide, 2013. Herbiguide. http://www.herbiguide.com.au/

Hoste I; Verloove F, 2009. Mediterranean container plants and their stowaways: a potential source of invasive plant species. Science facing Aliens Conference. Proceedings of a scientific meeting on Invasive Alien Species. Brussels, 11 May 2009. http://alienplantsbelgium.be/sites/alienplantsbelgium.be/files/Science%20Facing%20Aliens%20reprint%20IH-FV.pdf

HYPPA, 2013. Unité de Malherbologie & Agronomie. HYPPA.

INRA, 2013. Weed Science and Agronomy. Dijon, France: INRA. http://www2.dijon.inra.fr/hyppa/hyppa-a/eroci_ah.htm

Kew Royal Botanic Gardens, 2013. Seed Information Database. London, UK: Kew Royal Botanic Gardens

Kool A; Bengtson A; Thulin M, 2007. Polyphyly of Polycarpon (Caryophyllaceae) inferred from DNA sequence data. Taxon, 56(3):775-782.

Kurtto A; Lahti T, 1987. Checklist of the vascular plants of Finland. (Suomen putkilokasvien luettelo) Pamphlet of the Botanical Museum, University of Helsinki, 11:1-163.

Lunt ID, 1997. Germinable soil seed banks of anthropogenic native grasslands and grassy forest remnants in temperate south-eastern Australia. Plant Ecology, 130(1):21-34.

Mabberley DJ, 1997. The Plant Book: A Portable Dictionary of the Vascular Plants. 2nd edition. Cambridge, UK: Cambridge University Press.

Marañón T, 1998. Soil seed bank and community dynamics in an annual-dominated Mediterranean salt-marsh. Journal of Vegetation Science, 9(3):371-378.

McIntyre S; Lavorel S; Tremont RM, 1995. Plant life-history attributes: their relationship to disturbance response in herbaceous vegetation. Journal of Ecology (Oxford), 83(1):31-44.

Missouri Botanical Garden, 2013. Tropicos database. St Louis, USA: Missouri Botanical Garden. http://www.tropicos.org/

Mito T; Uesugi T, 2004. Invasive alien species in Japan: the status quo and the new regulation for prevention of their adverse effects. Global Environmental Research, 8:171-191.

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Links to Websites

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WebsiteURLComment
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.
Global register of Introduced and Invasive species (GRIIS)http://griis.org/Data source for updated system data added to species habitat list.

Contributors

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15/11/13: Original text by:

Ian Popay, consultant, New Zealand, with the support of Landcare Research.

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