Invasive Species Compendium

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Halophila stipulacea
(halophila seagrass)

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Datasheet

Halophila stipulacea (halophila seagrass)

Summary

  • Last modified
  • 16 December 2019
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Halophila stipulacea
  • Preferred Common Name
  • halophila seagrass
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Monocotyledonae
  • Summary of Invasiveness
  • H. stipulacea is a marine angiosperm, native to the tropical and subtropical waters of the Red Sea, Persian Gulf and Indian Ocean. It has spread to the Mediterranean and Caribbean seas. Fishing boats are the pr...

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Pictures

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PictureTitleCaptionCopyright
Halophila stipulacea (halophila seagrass); habit. South Sinai, Egypt. September 2013.
TitleHabit
CaptionHalophila stipulacea (halophila seagrass); habit. South Sinai, Egypt. September 2013.
Copyright©Tim Sheerman-Chase-2013/via flickr - CC BY 2.0
Halophila stipulacea (halophila seagrass); habit. South Sinai, Egypt. September 2013.
HabitHalophila stipulacea (halophila seagrass); habit. South Sinai, Egypt. September 2013.©Tim Sheerman-Chase-2013/via flickr - CC BY 2.0
Halophila stipulacea (halophila seagrass); habit. Caneel Bay, St. John US Virgin Islands. 2012.
TitleHabit
CaptionHalophila stipulacea (halophila seagrass); habit. Caneel Bay, St. John US Virgin Islands. 2012.
CopyrightPublic Domain - Released by National Oceanic & Atmospheric Administration (NOAA)/via flickr - CC0
Halophila stipulacea (halophila seagrass); habit. Caneel Bay, St. John US Virgin Islands. 2012.
HabitHalophila stipulacea (halophila seagrass); habit. Caneel Bay, St. John US Virgin Islands. 2012.Public Domain - Released by National Oceanic & Atmospheric Administration (NOAA)/via flickr - CC0
Halophila stipulacea (halophila seagrass); habit. (Artwork, from 'Symbolae physicae seu Icones Adhuc Inditeae : coroprum naturalium novorum aut minus cognitorum. Plate 5'. 1900.)
TitleHabit
CaptionHalophila stipulacea (halophila seagrass); habit. (Artwork, from 'Symbolae physicae seu Icones Adhuc Inditeae : coroprum naturalium novorum aut minus cognitorum. Plate 5'. 1900.)
CopyrightPublic Domain - From F.G. Hemprich & C.G. Ehrenberbg/''Symbolae physicae seu Icones Adhuc Inditeae : coroprum naturalium novorum aut minus cognitorum.' 1900.
Halophila stipulacea (halophila seagrass); habit. (Artwork, from 'Symbolae physicae seu Icones Adhuc Inditeae : coroprum naturalium novorum aut minus cognitorum. Plate 5'. 1900.)
HabitHalophila stipulacea (halophila seagrass); habit. (Artwork, from 'Symbolae physicae seu Icones Adhuc Inditeae : coroprum naturalium novorum aut minus cognitorum. Plate 5'. 1900.)Public Domain - From F.G. Hemprich & C.G. Ehrenberbg/''Symbolae physicae seu Icones Adhuc Inditeae : coroprum naturalium novorum aut minus cognitorum.' 1900.

Identity

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Preferred Scientific Name

  • Halophila stipulacea (Forssk.) Asch.

Preferred Common Name

  • halophila seagrass

Other Scientific Names

  • Thalassia stipulacea (Forssk.) K.D.Koenig
  • Zostera stipulacea Forssk.

International Common Names

  • English: broadleaf seagrass

Summary of Invasiveness

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H. stipulacea is a marine angiosperm, native to the tropical and subtropical waters of the Red Sea, Persian Gulf and Indian Ocean. It has spread to the Mediterranean and Caribbean seas. Fishing boats are the proposed pathway for the introduction of this species into the Mediterranean Sea, via the Suez Canal. A vector for transportation to the Caribbean is unknown, although recreational boats from the Mediterranean Sea have been proposed. Its environmental impacts include a reduction in native biodiversity and modification of successional patterns. It is included on a list of the 100 of the worst aliens species in Europe.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Monocotyledonae
  •                     Order: Hydrocharitales
  •                         Family: Hydrocharitaceae
  •                             Genus: Halophila
  •                                 Species: Halophila stipulacea

Notes on Taxonomy and Nomenclature

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The genus Halophila is comprised of 11 species, several of which have similar morphological features and habitat requirements, including the circumglobal seagrass H. decipiens.

Description

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H. stipulacea is a tropical, euryhaline marine angiosperm in Hydrocharitaceae, a family containing many aquatic plant species. Plant blades are elliptic, oblong and pale to dark green in colour with a length of 2-6 cm (Den Hartog, 1970; Willette and Ambrose, 2009). Pairs of blades extend from each rhizome node on petioles, and are covered at the base by folded and elliptic leaf scales 2-10 mm wide and 6-18 mm long. Blade margins, particularly at the apex, are serrated. Cross veins extend from the midrib to the intramarginal nerve at a 30° to 60° angle. Rhizomes are 0.5-2 cm in diameter, creeping and branched with a single root present at each node. Internodes have a length of 7-50 mm. Plant features are conservative between the native and invaded ocean basins, whereas blade and rhizome characteristics vary by depth (Den Hartog, 1970, Ruiz and Ballantine, 2004, Gambi et al., 2009, Willette and Ambrose, 2009).

Plant Type

Top of page Aquatic
Grass / sedge
Perennial
Vegetatively propagated

Distribution

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H. stipulacea is native to the tropical and subtropical waters of the Red Sea, Persian Gulf and Indian Ocean (DAISIE, 2015). It has spread, and is considered invasive, in the Mediterranean and Caribbean seas (GISD, 2015). In Egypt, it has been recorded in the Red Sea, where it is native, and also along its Mediterranean coast, where it is introduced (Gab-Alla, 2001; El-Hady et al., 2012).

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.

Last updated: 10 Jan 2020
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

ComorosPresentNativeGreen and Short (2003)
EgyptPresentNativeLipkin (1979); Gab-Alla (2001); El-Hady et al. (2012); CABI (Undated); Native in the Red Sea but introduced in the Mediterranean Sea
EritreaPresentNativeDen Hartog (1970)Massawa
KenyaPresentNativeDen Hartog (1970)Nyali Beach, Mombasa; Mokowe, Lamu; Gazi Bay
LibyaPresentIntroducedInvasiveSghaier et al. (2011)Al-wahesh Lagoon, Tobruk Bay
MadagascarPresentNativeDen Hartog (1970)Nossi Be; Fort Dauphin
MauritiusPresentNativeDen Hartog (1970)Grand River; Isle d’Ambre; Grand Bay
MozambiquePresentNativeGBIF (2014)
SeychellesPresentNativeCABI (Undated)La Rue Beach, Mahe Island; Original citation: Allem (1984)
TanzaniaPresentNativeGBIF (2014)
-Zanzibar IslandPresentIntroducedKamermans et al. (2002)Kiwengwa; Dongwe
TunisiaPresentIntroducedSghaier et al. (2011)Marina Cap Monastir; Kerkennah Island

Asia

BahrainPresentNativeDen Hartog (1970)Ras al Bar; Nuwaidrat
IndiaPresentNativeCABI (Undated a)Present based on regional distribution
-Tamil NaduPresent, WidespreadNativeParthasarathy et al. (1991)
IsraelPresentNativeMalm (2006); Schwarz and Hellblom (2002)Gulf of Aqaba
JordanPresentNativeWahbeh (1984); Abu-Hilal et al. (1988)Gulf of Aqaba
LebanonPresentIntroducedCABI (Undated); Lakkis and Novel-Lakkis (2007)Sidon; Beirut; Original citation: Lipkin (1975a)
QatarPresentNativeDen Hartog (1970)Doha
Saudi ArabiaPresentNativePrice and Cole (1992); Kenworthy et al. (1993)Mugeirma; Dawhat ad Dafi; multiple locations along Red Sea
TurkeyPresentIntroducedTurkozan and Durmus (2000); Ates et al. (2008); Meric et al. (2008)Fethiye Beach; Gumuldur
United Arab EmiratesPresentNativePhilips et al. (2002)Marawah; Jebel Dhanna; Dohat An Nakhla; Abu Dhabi Islands

Europe

AlbaniaPresentIntroducedSghaier et al. (2011)Karavasta Lagoon
CyprusPresent, WidespreadIntroduced1869InvasiveCABI (Undated)Original citation: Lipkin (1975b)
GreecePresent, WidespreadIntroduced1869InvasiveCABI (Undated); Malea (1994)Cape Mapatan; Aigina; Syros; Kythnos; Antikyra; Chios Island; Original citation: Lipkin (1975b)
ItalyPresentIntroducedInvasiveGambi et al. (2009); Procaccini et al. (1999)Palinuro harbour
MaltaPresentIntroducedInvasiveCABI (Undated)Marsaxlokk harbour; Original citation: Lipkin (1975a)

North America

ArubaPresentIntroduced2013InvasiveWillette et al. (2014)San Nicolas
CuraçaoPresentIntroduced2012InvasiveWillette et al. (2014)St. Joris Bay
DominicaPresentIntroduced2007InvasiveWillette et al. (2014); Willette and Ambrose (2009)Majority of west coastline
GrenadaPresentIntroduced2002InvasiveRuiz and Ballentine (2004)Flamingo Bay
GuadeloupePresent, WidespreadIntroduced2010InvasiveKerninon (2012)
MartiniquePresent, WidespreadIntroduced2008InvasiveWillette et al. (2014)
Netherlands AntillesPresentIntroduced2012InvasiveWillette et al. (2014)St. Eustatius
Saint LuciaPresentIntroduced2008InvasiveWillette and Ambrose (2009)Present in Anse La Raye, Marigot and Labrelotte Bays. Threat to native seagrasses
Saint Vincent and the GrenadinesPresent, WidespreadIntroduced2011InvasiveWillette et al. (2014)
U.S. Virgin IslandsPresentIntroduced2012InvasiveWillette et al. (2014)Widespread on St. John; localized on St. Thomas

South America

VenezuelaPresent, LocalizedIntroduced2013InvasiveVera et al. (2014)Puerto Azul; Playa Manza

History of Introduction and Spread

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H. stipulacea has spread to the Mediterranean and Caribbean seas, likely due to transportation by commercial and recreational shipping (GISD, 2015).

Mediterranean invasion – The opening and utilization of the Suez Canal beginning in 1869 aided the expansion of H. stipulacea to the Mediterranean Sea (Lipkin, 1975a). The seagrass was first found in Rhodes in 1984 by Johann Nemetz. Since 1987, H. stipulacea has been reported in the Tyrrhenian Sea (Gambi et al., 2009), Sicily (Bilotti and Abdelahad, 1990), Greece (Van der Velte and Den Hartog, 1992), Albania (Kashta and Pizzuto, 1995), Turkey (Alpinar, 1987), and most recently, Tunisia and Libya (Sghaier et al., 2011).

Caribbean invasion – H. stipulacea was first reported in the Caribbean in Flamingo Bay, Grenada in 2002 (Ruiz and Ballantine, 2004). In 2007, it was reported in Dominica (Willette and Ambrose, 2009) and has since been widely recorded throughout the Caribbean (Willette and Ambrose, 2009; Debrot et al., 2012; Kerninon, 2012). Most recently, it was reported along the coast of Venezuela (Vera et al., 2014).

Risk of Introduction

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No deliberate introductions of H. stipulacea to the Mediterranean or Caribbean Seas have been reported, however, dispersal potential may be high. Introduction of H. stipulacea to the Mediterranean from the Red Sea may have been via fishing boat traffic between basins (Lipkin, 1975a) and introduction to the Caribbean has been suggested to have occurred via recreational boat traffic between the Mediterranean and Caribbean Sea (Ruiz and Ballantine, 2004).

Habitat

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H. stipulacea is a marine plant that grows in sublittoral sediments on sand, mud and coral rubble (Galil, 2006).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Littoral
Coastal areas Principal habitat Harmful (pest or invasive)
Coastal areas Principal habitat Natural
Mud flats Principal habitat Harmful (pest or invasive)
Mud flats Principal habitat Natural
Intertidal zone Secondary/tolerated habitat Harmful (pest or invasive)
Intertidal zone Secondary/tolerated habitat Natural
Brackish
Estuaries Secondary/tolerated habitat Harmful (pest or invasive)
Estuaries Secondary/tolerated habitat Natural
Lagoons Principal habitat Harmful (pest or invasive)
Lagoons Principal habitat Natural
Marine
Coral reefs Principal habitat Harmful (pest or invasive)

Biology and Ecology

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Genetics

The chromosome number for H. stipulacea is 2n = 18 (Den Hartog, 1987).

Reproductive Biology

Asexual reproduction occurs via fragmentation and the dispersal of propagules on water currents.

Physiology and Phenology

Phenology of H. stipulacea is not well known in its invasive range. Flowers are rarely observed, but both flowers and fertile fruit-bearing plants were documented along Chios Island (Greece) in 2012 (Gerakaris and Tsiamis, 2015). Lipkin (1975b) collected both fruits and flowers in Rhodes in September, earlier in the year than fruits and flowers are typically observed in the northern Red Sea. Neither flowers nor fruit have yet been reported in the Caribbean Sea.

Climate

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ClimateStatusDescriptionRemark
A - Tropical/Megathermal climate Preferred Average temp. of coolest month > 18°C, > 1500mm precipitation annually
B - Dry (arid and semi-arid) Preferred < 860mm precipitation annually
C - Temperate/Mesothermal climate Preferred Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Depth (m b.s.l.) 2-45 0-147

Means of Movement and Dispersal

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Natural Dispersal

Fragmentation of plants creates viable propagules that settle in adjacent habitats or disperse via water currents. Sexually produced seeds are deposited in sediments where they are produced and may be distributed through natural disturbances (storms).

Accidental Introduction

Fragmentation of plants may be caused by and transported in fishing nets, fishing traps and boat anchors. Fishing boats are the proposed pathway for the introduction of this species into the Mediterranean Sea, via the Suez Canal (Lipkin, 1975a). A vector for transportation to the Caribbean is unknown, although recreational boats from the Mediterranean Sea have been proposed (Ruiz and Ballantine, 2004). Within-basin dispersal may occur by local recreational and fishing boat activities as well as storms that liberate and disperse seagrass propagules. 

Pathway Causes

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CauseNotesLong DistanceLocalReferences
FisheriesAnecdotal and documented observations of H. stipulacea in fishing nets and traps that are transported to new locations (e.g. Caribbean and Mediterranean seas) Yes Yes Ruiz and Ballantine, 2004; Willette et al., 2014
Interbasin transfersAccidental Yes Lipkin, 1975a

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Water Yes
Ship ballast water and sediment Yes Yes Ruiz and Ballantine, 2004

Impact Summary

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CategoryImpact
Environment (generally) Negative

Environmental Impact

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According to Galil (2006), H. stipulacea is inducing changes to sublittoral communities in the Mediterranean and out-competing native Mediterranean seagrasses. It is listed as an invasive species in the Mediterranean Sea (Bourdouresque and Verlaque, 2002) and is also included on a list of the 100 worst aliens species in Europe (DAISIE, 2015). 

Elsewhere, the environmental impacts of H. stipulacea are largely unknown. However, extirpation of congener Halophila decipiens from the island of Dominica (Caribbean) was reported, where invasive H. stipulacea now occurs in habitats where H. decipiens previously existed (Steiner and Willette, 2015a). Also in Dominica, there is evidence that H. stipulacea has caused significant changes to native seagrass meadows, primarily by replacing native Syringodium filiforme, as well as partially overgrowing coral reefs (Steiner and Willette, 2015b). Changes in fish composition have also been reported in Dominica and the US Virgin Islands (Willette and Ambrose, 2012; Olinger et al., 2017).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Highly adaptable to different environments
  • Pioneering in disturbed areas
  • Tolerant of shade
  • Fast growing
  • Reproduces asexually
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Modification of successional patterns
  • Reduced native biodiversity
Impact mechanisms
  • Competition (unspecified)
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally

Similarities to Other Species/Conditions

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Despite differences in morphology, H. stipulacea may have been confused with congener H. decipiens in the Caribbean Sea, which has overlapping ecological requirements.

References

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Abu-Hilal A, Badran M, de Vaugelas J, 1988. Distribution of trace elements in Callichirus laurae burrows and nearby sediments in the Gulf of Aqabat, Jordan (Red Sea). Marine Environmental Research, 25, 233-248.

Aleem AA, 1984. Distribution and ecology of seagrass communities in the Western Indian Ocean. Deep Sea Research Part A. Oceanographic Research Papers, 31(6-8), 919-933.

Alpinar K, 1987. A new record for Turkish flora Halophila stipulacea. Istanbul Universitesi Eczacilik Fakultesi Mecmuasi, 23, 83-84.

Ates A, Özdilek S, Özcan T, Kontas T, 2008. On the presence of the brachyuran crab Macropodia tenuirostris (Crustacea: Decapoda: Majidae) on the Levantine coast of Turkey. Marine Biodiversity Records, 1(e79), doi: 10.1017/S1755267207008020

Biliotti CN, Abdelahad N, 1990. Halophila stipulacea (Forrsk.) Aschers. (Hydrochoritaceae): New Species for Italy. (Halophila stipulacea (Forssk.) Aschers. (Hydrocharitaceae): espèce nouvelle pour l’Italie). In: Posidonia Newsletter , 3. 23-26.

Boudouresque, C. F., Verlaque, M., 2002. Biological pollution in the Mediterranean Sea: invasive versus introduced macrophytes. Marine Pollution Bulletin, 44(1), 32-38. doi: 10.1016/S0025-326X(01)00150-3

DAISIE, 2015. Delivering Alien Invasive Species Inventories for Europe. In: European Invasive Alien Species Gateway . www.europe-aliens.org/default.do

Debrot AO, Hylkema A, Vogelaar W, Meesters HWG, Engel MS, Leon R, Prud'homme van Reine WF, Nagelkerken I, 2012. Baseline surveys of Lac Bay benthic and fish communities, Bonaire. In: IMARES Report C129/12 , Netherlands: Institute for Marine Resource Ecosystem Studies.

Den Hartog C, 1970. The sea-grasses of the world, London, UK: North-Holland.

Den Hartog C, Hennen J, Noten TMPA, van Wijk RJ, 1987. Chromosome numbers of the European seagrasses. Plant Systematics and Evolution, 156(1-2), 55-59.

El-Hady, H. H. A., Hamed, E. R., Shehata, A. N., 2012. Molecular identification, antimicrobial and antioxidant activities of the tropical seagrass Halophila stipulacea grown in El-Bardawil lake, Egypt. Australian Journal of Basic and Applied Sciences, 6(12), 474-481. http://www.ajbasweb.com/ajbas/2012/Nov%202012/474-481.pdf

Gab-Alla AAFA, 2001. Ecological status of seagrass communities in Sharm El-Moyia Bay (Gulf of Aqaba, Red Sea) after oil pollution in 1999. Journal of King Abdulaziz University - Marine Sciences, 12, 231-239.

Gallil BS, 2006. Halophila stipulacea. DAISIE.http://www.europe-aliens.org/pdf/Halophila_stipulacea.pdf

Gambi M, Barbieri F, Bianchi C, 2009. New record of the alien seagrass Halophila stipulacea (Hydrocharitaceae) in the western Mediterranean: A further clue to changing Mediterranean Sea biogeography. Marine Biodiversity Records, 2(e84), doi: 10.1017/S175526720900058X

GBIF, 2014. Global Biodiversity Information Facility. http://www.gbif.org/species

Gerakaris, V., Tsiamis, K., 2015. Sexual reproduction of the Lessepsian seagrass Halophila stipulacea in the Mediterranean Sea. Botanica Marina, 58(1), 51-53. http://www.degruyter.com/view/j/botm doi: 10.1515/bot-2014-0091

GISD, 2015. Global Invasive Species Database (GISD). http://www.iucngisd.org/gisd/

Green EP, Short FT, 2003. World Atlas of Seagrasses, Berkeley, USA: UNEP-WCMC and University of California Press.

Jacobs RPWM, Dicks B, Seagrasses in the Zeit Bay area and at Ras Ghârib (Egyptian Red Sea coast). Aquatic Botany, 23(1), 137-147.

Kamermans, P., Hemminga, M. A., Tack, J. F., Mateo, M. A., Marbà, N., Mtolera, M., Stapel, J., Verheyden, A., Daele, T. van, 2002. Groundwater effects on diversity and abundance of lagoonal seagrasses in Kenya and on Zanzibar Island (East Africa). Marine Ecology, Progress Series, 231, 75-83. doi: 10.3354/meps231075

Kashta L, Pizzuto F, 1995. On the presence of Halophila stipulacea (Forskal) Ascherson on the coasts of Albania. (Sulla presenza di Halophila stipulacea (Forskal) Ascherson nelle coste dell'Albania). Bollettino delle Sedute della Accademia Gioenia di Scienze Naturali in Catania, 28, 161-166.

Kenworthy WJ, Durako MJ, Fatemy SMR, Valavi H, Thayer GW, 1993. Ecology of seagrasses in northeastern Saudi Arabia one year after the Gulf War oil spill. Marine Pollution Bulletin, 27, 213-222.

Kerninon F, 2012. First actions in setting up an overseas seagrass monitoring network. (Première actions de mis en place d’un réseau d’observation des herbiers de l’Outre-mer). Universite de Bretagne Occidentale. 136 pp. http://ifrecor-doc.fr/items/show/1478

Lakkis S, Novel-Lakkis V, 2007. Diversity and distribution of macrophytes along the coast of Lebanon (Levantine Basin, Eastern Mediterranean). Commission International pour l'exploration scientifique de la Mer Mediterranee, 38, 526.

Lipkin Y, 1975a. Halophila stipulacea, a review of a successful immigration. Aquatic Botany, 1, 203-215.

Lipkin Y, 1975b. Halophila stipulacea in Cyprus and Rhodes, 1967-1970. Aquatic Botany, 1, 309-318.

Lipkin Y, 1979. Quantitative aspects of seagrass communities, particularly of those dominated by Halophila stipulacea, in Sinai (Northern Red Sea). Aquatic Botany, 7, 119-128.

Malea P, 1994. Seasonal variation and local distribution of metals in the seagrass Halophila stipulacea (Forsk.) Aschers. in the Antikyra Gulf, Greece. Environmental Pollution, 85, 77-85.

Malm T, 2006. Reproduction and recruitment of the seagrass Halophila stipulacea. Aquatic Botany, 82(4), 345-349.

Meric E, Yokes MB, Avsar N, 2008. Amphisorus hemprichii Ehrenberg (Rhizopoda, foraminifera) along the Antalya coast. Micropaleontology, 54(3), 277-292.

Olinger LK, Heidmann SL, Durdall AN, Howe C, Ramseyer T, Thomas SG, Lasseigne DN, Brown EJ, Cassell JS, Donihe MM, Duffing Romero MD, Duke MA, Green D, Hillbrand P, Wilson Grimes KR, Nemeth RS, Smith TB, Brandt M, 2017. Altered juvenile fish communities associated with invasive Halophila stipulacea seagrass habitats in the U.S. Virgin Islands. PLOS ONE, 12(11), e0188386. doi: 10.1371/journal.pone.0188386

Parthasarathy N, Ravikumar K, Ganesan R, Ramanurthy K, 1991. Distribution of seagrasses along the coast of Tamil Nadu, Southern India. Aquatic Botany, 40, 145-153.

Philips RC, Loughland RA, Youssef A, 2002. Seagrasses of Abu Dhabi. UAE Tribulus, 12(1), 20-23.

Price ARG, Cole SL, 1992. Aspects of seagrass ecology along the western Arabian Gulf coast. Hydrobiologica, 234, 129-141.

Procaccini G, Acunto S, Fama P, Maltagliati F, 1999. Structural, morphological and genetic variability in Halophila stipulacea (Hydrocharitaceae) populations in the western Mediterranean. Marine Biology, 135(1), 181-189.

Ruiz H, Ballentine DL, 2004. Occurrence of the seagrass Halophila stipulacea in the tropical west Atlantic. Bulletin of Marine Science, 75(1), 131-135.

Schwarz AM, Hellblom F, 2002. The photosynthetic light response of Halophila stipulacea growing along a depth gradient in the Gulf of Aqaba, the Red Sea. Aquatic Botany, 74, 263-272.

Sghaier YR, Zakhama-Sraieb R, Benamer I, Charfi-Cheikhrouha F, 2011. Occurrence of the seagrass Halophila stipulacea (Hydrocharitaceae) in the southern Mediterranean Sea. Botanica Marina, 54, 575-582.

Steiner SCC, Willette DA, 2015a. The expansion of Halophila stipulacea (Hydrocharitaceae, Angiospermae) is changing the seagrass landscape in the Commonwealth of Dominica, Lesser Antilles. Caribbean Naturalist, 22, 1-19.

Steiner SCC, Willette DA, 2015b. Dimming sand halos in Dominica and the expansion of the invasive seagrass Halophila stipulacea. Reef Encounter, 30, 43-44.

Turkozan O, Durmus SH, 2000. A feeding ground for juvenile green turtles, Chelonia mydas, on the western coast of Turkey. British Herpetological Society Bulletin, 71, 1-5.

Van der Velte G, den Hartog C, 1992. Continuing range extension of Halophila stipulacea (Forssk.) Aschers. (Hydrocharitaceae) in the Mediterranean - Now found at Kefallinia and Ithaki (Ionian Sea). Acta Botanica Neerlandica, 41(3), 345-348.

Vera B, Collado-Vides L, Moreno C, van Tussenbroek BI, 2014. Halophila stipulacea (Hydrocharitaceae): A recent introduction to the continental waters of Venezuela. Caribbean Journal of Science, 48(1), 66-70.

Wahbeh MI, 1984. The growth and production of the leaves of the seagrass Halophila stipulacea (Forsk.) Aschers. from Aqaba, Jordan. Aquatic Botany, 20, 33-41.

Willette DA, Ambrose RF, 2009. The distribution and expansion of the invasive seagrass Halophila stipulacea in Dominica, West Indies, with a preliminary report from St. Lucia. Aquatic Botany, 91(3):137-142. http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6T4F-4W38RMT-1&_user=10&_coverDate=10%2F31%2F2009&_rdoc=4&_fmt=high&_orig=browse&_srch=doc-info(%23toc%234973%232009%23999089996%231461076%23FLA%23display%23Volume)&_cdi=4973&_sort=d&_docanchor=&_ct=22&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=8ee3881fee02179736263184e13b817f

Willette, D. A., Ambrose, R. F., 2012. Effects of the invasive seagrass Halophila stipulacea on the native seagrass, Syringodium filiforme, and associated fish and epibiota communities in the Eastern Caribbean. Aquatic Botany, 103, 74-82. http://www.sciencedirect.com/science/article/pii/S0304377012001209 doi: 10.1016/j.aquabot.2012.06.007

Willette, D. A., Chalifour, J., Debrot, A. O. D., Engel, M. S., Miller, J., Oxenford, H. A., Short, F. T., Steiner, S. C. C., Védie, F., 2014. Continued expansion of the trans-Atlantic invasive marine angiosperm Halophila stipulacea in the Eastern Caribbean. Aquatic Botany, 112, 98-102. http://www.sciencedirect.com/science/article/pii/S0304377013001289 doi: 10.1016/j.aquabot.2013.10.001

Distribution References

Abu-Hilal A, Badran M, de Vaugelas J, 1988. Distribution of trace elements in Callichirus laurae burrows and nearby sediments in the Gulf of Aqabat, Jordan (Red Sea). In: Marine Environmental Research, 25 233-248.

Ates A, Özdilek S, Özcan T, Kontas T, 2008. On the presence of the brachyuran crab Macropodia tenuirostris (Crustacea: Decapoda: Majidae) on the Levantine coast of Turkey. In: Marine Biodiversity Records, 1 (e79) DOI:10.1017/S1755267207008020

CABI, Undated. Compendium record. Wallingford, UK: CABI

CABI, Undated a. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI

CABI, Undated b. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

Den Hartog C, 1970. The sea-grasses of the world., London, UK: North-Holland.

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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.
Invasive Seagrasshttp://www.invasiveseagrass.org/Citizen science website to track the spread of Halophila stipulacea

Organizations

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USA: SeagrassNet, University of New Hampshire, www.seagrassnet.org

Australia: Seagrass-Watch, James Cook University, www.seagrasswatch.org

Contributors

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24/03/15 Original text by:

Demian A Willette, University of California, Los Angeles, California, USA

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