Overview

Comprehensive Description

Halodule wrightii is a "seagrass" that may form carpet-like beds in warm, shallow waters from the southeastern United States to South America (seagrasses superficially resemble grasses, but are not technically grasses since they are not in the family Poaceae) (Haynes 2000).

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Distribution

Range Description

Halodule wrightii has a disjunct global and predominantly tropical distribution. The main part of its range is in the Atlantic: the western tropical Atlantic from northern Florida (USA) to Venezuela including the Gulf of Mexico and the Caribbean Sea; also in Bermuda and North Carolina (USA). In the southern Atlantic, it is present on the coast of Brazil. In the eastern Atlantic, it occurs from southern Morocco on the coast of Africa and the Canary Islands, to the northern part of Angola.

Halodule wrightii is found in the eastern tropical Pacific from the Gulf of California to the Gulf of Panama. In the Indian Ocean it is found from the northern extent of the Bay of Bengal to along the Coromandel Coast as well as Oman. Also from southern Somalia to the north part of South Africa including the Mozambique Channel, Mauritius and Madagascar.
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National Distribution

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

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Global Range: Native to coastal Mexico and Caribbean Sea. Presumed extirpated from the Salton Sea, Calif. (Hickman, 1993).

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Ala., Fla., La., Miss., N.C. , Tex.; e Mexico; West Indies; Central America (Guatemala, Belize, Nicaragua); South America (Venezuela).
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Halodule wrightii is the most common seagrass species in Brazil. This species approaches its southern distributional limit along the Rio de Janeiro state coast (Creed,1997). The northern limit of its range along the Atlantic coast of North America is North Carolina (Ferguson et al.1993).

The range of Halodule wrightii includes Alabama, Florida, Louisiana, Mississippi, and North Carolina (U.S.A.); eastern Mexico; the West Indies; Central America (Belize, Guatemala, Nicaragua); and South America (Venezuela) (Haynes 2000).

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Physical Description

Morphology

Description

Rhizomes: internodes 0.5----4 cm; scales 4----10 mm. Leaves dark red-brown; sheath 1.5--6 cm ´ 1--2 mm; blade 5--20 cm ´ 0.3--1.5 mm, apex notched or with very prominent, acute median tooth; veins ending in teeth or not. Inflorescences solitary. Staminate flowers: peduncles 10--25 mm; distal anthers ca. 0.5 mm higher than the proximal. Pistillate flowers: styles lateral to terminal. Fruits ovoid to globose, 1.5--2 ´ 1.2--1.8 mm.
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Diagnostic Description

Synonym

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Type Information

Isotype for Halodule wrightii Asch.
Catalog Number: US 43433
Collection: Smithsonian Institution, National Museum of Natural History, Department of Botany
Verification Degree: Card file verified by examination of alleged type specimen
Preparation: Pressed specimen
Collector(s): C. Wright
Locality: Greater Antilles, Cuba, West Indies
  • Isotype: Ascherson, P. F. 1868. Sitzungsber. Ges. Naturf. Freunde Berlin. 1868: 19.
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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
Halodule wrightii is typically found on sandy to muddy bottoms and can be found in mixed seagrass species beds. It is highly tolerant to a range of environmental conditions including wide ranging salinity (hypersaline), high temperatures, turbidity, and eutrophication (Zieman 1982, UNESCO 1998, Hemminga and Duarte 2000, Green and Short 2003, Larkum et al. 2006). Optimum temperatures for H. wrightii range between 20-30°C (Phillips 1960).

This species is ephemeral with rapid turn-over and high seed set and forming effective seedbanks, well adapted to high levels of disturbance. It is a pioneer species in Mozambique in exposed sandy areas close to the coastline. It is the dominant species in Texas (USA). It is established along both the eastern and western margins of the lagoon in Tamaulipas (Mexico). In Veracruz (Mexico), it is found in the shallower areas where it tolerates changes in temperature and salinity. In the Caribbean, it is found growing on sand and mud from the intertidal zone to five m. It is the most widely distributed seagrass in Brazil. In South America, it is associated with shallow habitats without much freshwater input, such as reefs, algal beds, coastal lagoons, rocky shores, sand beaches, and unvegetated soft-bottom areas and nearby mangroves (Green and Short 2003).

After a complete destruction of seabeds, a rapid recovery was observed with early recovery characterized by small patches suggesting recovery through fragments. Studies show that fragments stay viable for up to four weeks in the spring months and up to two weeks during the autumn months. High viability of fragments suggests a high dispersal distance (Hall et al. 2006).

This is usually an early colonizing species yet studies in Florida Bay shows that with increased nutrient levels, Halodule wrightii becomes the dominant species as it is able to out-compete Thalassia testudinum for light resources, suggesting that areas in Florida Bay with high nutrient availability will be dominated by Halodule wrightii while areas with low nutrients will be dominated by T. testudinum (Fourqurean et al. 1995).

Halodule wrightii can rapidly and densely recolonize denuded areas in warm months. Most bed maintenance and new shoot production probably occurs through rhizome elongation (Phillips 1960).

Systems
  • Marine
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Comments: Marine or saltwater-aquatic (e.g., Salton Sea [formerly] Expected in warm, coastal marine habitats in the Atlantic, Pacific, and Caribbean.

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Intertidal zone of marine waters with sandy or muddy substrates; -2--0m.
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Depth range based on 176 specimens in 1 taxon.
Water temperature and chemistry ranges based on 165 samples.

Environmental ranges
  Depth range (m): 0 - 89
  Temperature range (°C): 25.700 - 27.678
  Nitrate (umol/L): 0.161 - 1.248
  Salinity (PPS): 35.179 - 37.096
  Oxygen (ml/l): 4.603 - 4.746
  Phosphate (umol/l): 0.020 - 0.125
  Silicate (umol/l): 0.888 - 2.517

Graphical representation

Depth range (m): 0 - 89

Temperature range (°C): 25.700 - 27.678

Nitrate (umol/L): 0.161 - 1.248

Salinity (PPS): 35.179 - 37.096

Oxygen (ml/l): 4.603 - 4.746

Phosphate (umol/l): 0.020 - 0.125

Silicate (umol/l): 0.888 - 2.517
 
Note: this information has not been validated. Check this *note*. Your feedback is most welcome.

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Halodule wrightii is found in the intertidal zone of marine waters with sandy or muddy substrates at depth from 0 to 2 meters. Halodule wrightii occupies the shallowest waters in the Gulf of Mexico and is often exposed during low tides. (Haynes 2000)

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Associations

Seagrass beds may be disrupted by the activities of a variety of animals, including horseshoe crabs (Limulus polyphemus), Cownose Rays (Rhinoptera bonasus), and Southern Stingrays (Dasyatis sabina). One mode of foraging that has been reported for manatees (Trichechus manatus) involves the use of their forelimbs to uproot seagrass, leaving scattered roots and blades behind. These disturbances have generally been viewed as having a negative impact on seagrass beds. Hall et al. note, however, that if the fragments created by these animals are capable of settling and rooting elsewhere, the result may be the formation of new seagrass patches. (Hall et al. 2006)

Taplin et al. (2005) investigated interactions between H. wrightii and the macroalga Caulerpa prolifera. Their experiments indicated that the density and biomass of H. wrightii were negatively influenced by the presence of C. prolifera. Whether the nature of the interaction was a result of competition for space, nutrients or light could not be determined from their study. The performance of both species, however, differed between the two water depths studied, suggesting that the outcome of the interaction may be moderated in some way by light levels.

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Known predators

Halodule wrightii (Seagrass) is prey of:
Dasyatis sabina
Arius felis
Anas discors
dissolved organic carbon
sediment POC
Elasmopus levis
Lembos rectangularis
Acunmindeutopus naglei
Synchelidium
Ampithoe longimana
Cymadusa compta
Batea catharinensis
Listriella barnardi
Lysianopsis alba
Caprella penantis
Libinia dubia
Pinixia floridana
Neopanope texana
Paracerces caudata
Erichsionella
Edotia triloba
Alpheus normani
Hippolyte zostericola
Processa bermudiensis
Penaeus duoarum
Palaemonetes floridanus

Based on studies in:
USA: Florida (Estuarine)

This list may not be complete but is based on published studies.
  • Christian RR, Luczkovich JJ (1999) Organizing and understanding a winter’s seagrass foodweb network through effective trophic levels. Ecol Model 117:99–124
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Population Biology

Number of Occurrences

Note: For many non-migratory species, occurrences are roughly equivalent to populations.

Estimated Number of Occurrences: 81 to >300

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General Ecology

Ecology

In an assessment of light requirements of H. wrightii, Kenworthy and Fonseca (1996) found that the lower limit of depth distribution was controlled by light availability. Estimates of minimum light requirements ranged from about a quarter to a third of the light just beneath the water surface, much higher than light levels in the photic zone for many phytoplankton and macroalgae (typically just 1 to 5% of incident light).

Burd and Dunton (2001) used long-term data on the biomass of Halodule wrightii in the Upper Laguna Madre, Texas, to validate a model demonstrating the importance of underwater light intensity as a major abiotic factor regulating H. wrightii productivity.

Heck et al. (2006) assessed the individual and combined effects of removing large predators and enriching water column nutrients on Halodule wrightii meadows in Big Lagoon, Florida, U.S.A. To simulate the first-order effects of large predator reductions, the authors stocked enclosures with ~3 to 4 times natural densities of the omnivorous pinfish Lagodon rhomboides, the dominant
fish in local seagrass habitats, and supplemented nitrogen and phosphorus in the water column to nearly 3 times normal levels. Results showed both significant predator and nutrient effects, although there were fewer consumer effects and more negative nutrient effects on seagrasses than had been found in previous work, which had shown that mesograzers ameliorated the harmful effects of elevated nutrients on seagrasses. Epiphyte proliferation in nutrient enrichment treatments did not occur; thus, algal overgrowth could not explain the negative effects of nutrient loading on seagrass biomass. Instead, nutrient loading resulted in nitrogen-rich shoalgrass, and the authors suggest that this high-quality food stimulated pinfish herbivory, resulting in the decline of seagrass biomass in enrichment enclosures.

Armitage and Fourqurean (2006) transplanted H. wrightii sprigs into caged and uncaged plots in a Turtlegrass (Thalassia testudinum) bed near a patch reef. Nutrients (nitrogen and phosphorus) were added to half of the experimental plots. The authors recorded changes in seagrass shoot density, and after three months, measured above- and below-ground biomass and tissue
nutrient content of both Turtlegrass and H. wrightii. Herbivory immediately and strongly impacted H. wrightii. Within six days of transplantation, herbivory reduced the density of uncaged H. wrightii by over 80%, resulting in a decrease in above- and below ground biomass of nearly an order of magnitude. Turtlegrass shoot density and below-ground biomass were not affected by herbivory, but above-ground biomass and leaf surface area were higher within cages, suggesting that herbivory influenced both seagrass species, but that Turtlegrass was more resistant to herbivory pressure than was H. wrightii. Nutrient addition did not alter herbivory rates or the biomass of either species over the short-term duration of this study. In both species, nutrient addition had little effect on the tissue nutrient content of seagrass leaves, and the nitrogen-to-phosphorus ratio was near the 30:1 threshold ratio that suggests a balance in supply, indicating that neither of these elements appeared to be limiting growth. The authors note that the different impacts of grazing on these two seagrass species suggest that herbivory may be an important regulator of the distribution of multiple seagrass species near herbivore refuges like patch reefs in the Caribbean.

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Life History and Behavior

Cyclicity

Flowering/Fruiting

Flowering summer.
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Reproduction

Like most seagrasses, Halodule wrightii is highly clonal, extending and maintaining perennial beds through the growth of underground rhizomes (McGovern and Blankenhorn 2007 and references therein).

Hall et al. (2006) found that vegetative fragments of Halodule wrightii can settle and re-root. Settlement of uprooted vegetative fragments may be a viable recruitment mechanism in some areas, especially where sexual reproduction (via flowering anfd fruiting) is not occurring or is rare. Fragments of the fast growing H. wrightii have the ability to remain viable for periods of time that allow for dispersal over large distances (kilometers). Morris & Virnstein (2004) observed a rapid recovery of H. wrightii in the northern Indian River Lagoon, Florida (U.S.A.), in quiescent, shallow water. This recovery occurred after complete demise of the existing seagrass bed. The initial recovery occurred as small patches, which may be indicative of recruitment by H. wrightii fragments. Hall et al. found that in their experiments H. wrightii was more successful at recruitment via fragments in spring than in fall. This species grows fastest throughout the spring and summer months and enters a period of dormancy during the late fall and winter months. Spring fragments of H. wrightii appear to have the potential to travel much greater distances in 4 weeks than do fall fragments, which lose viability by the second week. In addition to viability, of course, the distances the fragments can travel depend also on factors such as wind and tidal currents. (Hall et al. 2006)

In some parts of the range of H. wrightii, flowering and fruiting are rare, but in other areas these are apparently regular events (McGovern and Blankenhorn 2007 and references therein). In North Carolina, flowering has been reported from May to August and over a wide range of salinity (12 to 34 ppt). Flowers occurred at water depths from exposed to 1.5 meters deep at low tide. Halodule wrightii is dioecious (i.e., individual plants are either male or female rather than both). (Ferguson et al. 1993)

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Growth

Development

In a study of Halodule wrightii in the Gulf of Mexico off Alabama (U.S.A.), the total biomass of H. wrightii generally increased through late summer, then began to decline. The proportion of that mass accounted for by roots and rhizomes generally declined from a high level in mid-spring as the proportion comprising leaves generally increased. The mass of below-ground structures was always greater than that of above-ground structures. (McGovern and Blankenhorn 2007)

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Evolution and Systematics

Evolution

Systematics and Taxonomy

For many years Halodule along the North American coast were considered to be distinct from H. wrightii and referred to as H. beaudettei. Based on studies of variation in leaf tip shape in the northern Gulf of Mexico, however, along, along with a failure to identify genetic (isozyme) differences between plants with the different morphologies supposedly representing distinct species, Haynes concluded that H. beaudettei should be treated as a synonym of H. wrightii (Haynes 2000 and references therein)

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Molecular Biology and Genetics

Molecular Biology

Barcode data: Halodule wrightii

The following is a representative barcode sequence, the centroid of all available sequences for this species.


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Statistics of barcoding coverage: Halodule wrightii

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 2
Specimens with Barcodes: 2
Species With Barcodes: 1
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2010

Assessor/s
Short, F.T., Carruthers, T.J.R., van Tussenbroek, B. & Zieman, J.

Reviewer/s
Livingstone, S., Harwell, H. & Carpenter, K.E.

Contributor/s

Justification
Halodule wrightii has a disjunct global distribution. The main part of its range is in the Atlantic, although it is also found in the eastern tropical Pacific and the Indian Ocean. It is the dominant species in Brazil and West Africa. Halodule wrightii is a widespread species and is locally abundant. The overall population trend for this species is stable, and possibly increasing in some parts of its range. It is highly tolerant to a range of environmental conditions, however, it is affected locally by coastal development and destructive anthropogenic activities. This species is listed as Least Concern.
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National NatureServe Conservation Status

United States

Rounded National Status Rank: N4 - Apparently Secure

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NatureServe Conservation Status

Rounded Global Status Rank: G5 - Secure

Reasons: Abundant in Caribbean, northward to North Carolina, and along the Pacific coast of Mexico and Central America.

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Population

Population
Halodule wrightii is a widespread species which is locally abundant. It is the dominant species in Brazil and West Africa. In other areas it may be found in mixed beds with other seagrass species. Its presence on the western side of Central America appears to be a migration through the Panama Canal and therefore its range appears to be expanding and there is potential for future range extension if it is introduced into other areas.

According to the Global Seagrass Trajectories Database, (Carruthers pers. comm. 2007) there are 64 published studies that monitored this species over time and of these 40 had no change, 15 decreased in coverage, and nine increased in coverage (all aerial extent, density, biomass, or cover). The overall population trend for this species is increasing (2% increase). Data from 2000 shows a 28% occurrence when sampling 188 stations at Big Bend, Florida. This is a 23% increase from the data reported by Iverson and Bittaker (1986). Despite the increase in occurrence, maximum depth range decreased from 10.6-8.3 m (Hale et al. 2004). Research by Hall et al. (1999) show a significant widespread decrease in the Florida Bay with a bay wide short shoot density off about 267.5 shoots/m² in 1984 to about 22.5 shoots/m² in 1994. Decrease in abundance was most likely cause by increased light attenuation due to primary die-off off Thalassia testudinum. Reduction can also be linked to a decrease in phosphorus availability caused by reduction in freshwater input (Hall et al. 1999). Global average maximum biomass is estimated to be 253.5 g dw/m² above ground (from 19 observations) and 193.3 g dw/m² below ground (from 12 observations) (Duarte and Chiscano 1999).

Population Trend
Increasing
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Threats

Major Threats
Halodule wrightii is a tolerant species to most disturbances. It replaces less tolerant species under conditions of habitat deterioration, eutrophication, and increased turbidity, and therefore general threats are not considerable except in localized situations. Localized threats include trawling activities, coastal development, habitat destruction and mechanical damage from anchoring and recreational and commercial boating.
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Management

Conservation Actions

Conservation Actions
Halodule wrightii is protected in numerous marine protected areas throughout its range.
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Wikipedia

Halodule wrightii

Halodule wrightii is a plant species native to seacoasts of some of the warmer oceans of the world. It has been reported from California, Texas, Florida, Louisiana, Mississippi, Alabama, North Carolina, Maryland, Yucatán, Quintana Roo, Tabasco, Costa Rica, Belize, Panamá, Cuba, Trinidad & Tobago, Venezuela, Brazil, Australia and Madagascar.[3][4][5][6][7][8][9][10][11]

Some publications cite US specimens by the synonym, Halodule beaudettei,[12][13] but the two names represent the same species.[3][14][15][16]

Halodule wrightii is an herb growing in salt-water marshes in intertidal regions, often submerged at high tide but emergent at low tide. It has flat leaves up to 20 cm long, dark reddish-brown, with a few teeth on the margins. Fruits are spherical to egg-shaped, about 2 mm across.[3][17][18][19][20]

References[edit]

  1. ^ Tropicos
  2. ^ The Plant List
  3. ^ a b c Flora of North America v 22
  4. ^ Hickman, J. C. 1993. The Jepson Manual: Higher Plants of California 1–1400. University of California Press, Berkeley.
  5. ^ Hammel, B. E. 2003. Cymodoceaceae. In: Manual de Plantas de Costa Rica, B.E. Hammel, M.H. Grayum, C. Herrera & N. Zamora (eds.). Monographs in systematic botany from the Missouri Botanical Garden 92: 456–457.
  6. ^ Novelo R., A. & A. L. H. 1994. 239. Cymodoceaeceae. 6: 15–16. In G. Davidse, M. Sousa Sánchez & A.O. Chater (eds.) Flora Mesoamericana. Universidad Nacional Autónoma de México, México, D. F.
  7. ^ Balick, M. J., M. H. Nee & D.E. Atha. 2000. Checklist of the vascular plants of Belize. Memoirs of The New York Botanical Garden 85: i–ix, 1–246.
  8. ^ Correll, D. S. & M. C. Johnston. 1970. Manual of the Vascular Plants of Texas i–xv, 1–1881. The University of Texas at Dallas, Richardson
  9. ^ Cowan, C. P. 1983. Flora de Tabasco. Listados Florísticos de México 1: 1–123.
  10. ^ Sousa Sánchez, M. & E. F. Cabrera Cano. 1983. Flora de Quintana Roo. Listados Florísticos de México 2: 1–100.
  11. ^ BONAP (Biota of North America Project) floristic synthesis, Halodule wrightii
  12. ^ Hartog, Cornelis den. 1964. Blumea 12: 303.
  13. ^ Hartog, Cornelis den. 1960. Pacific Naturalist 1(15): 4–5, f. 2a–c.
  14. ^ Phillips, R. C. 1967. On species of the seagrass, Halodule, in Florida. Bull. Mar. Sci. 17: 672--676.
  15. ^ McMmillan, C. 1991. Isozyme patterning in marine spermatophytes. In: L. Triest, ed. 1988+. Isozymes In Water Plants. Opera Botanica Belgica 1+ vols. Belgium, Meise. Vol. 4,: pp. 193--200.
  16. ^ photo of specimen of Halodule wrightii at Missouri Botanical Garden, collected in Tabasco
  17. ^ Novelo, A. & L. Ramos. 2005. Vegetación acuática. Cap. 5: 111–144. In J. Bueno, F Álvarez & S. Santiago, Biodiversidad del Estado de Tabasco. CONABIO-UNAM, México.
  18. ^ Godfrey, R. K. & J. W. Wooten. 1979. Aquatic and Wetland Plants of Southeastern United States Monocotyledons 1–712. The University of Georgia Press, Athens.
  19. ^ Ascherson, Paul Friedrich August. 1868. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1868: 19.
  20. ^ Ascherson, Paul Friedrich August. 1897. Die Natürlichen Pflanzenfamilien 2: 37.
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Notes

Comments

Halodule wrightii occupies the shallowest waters in the Gulf of Mexico. In fact, the plants are often exposed during low tides. 

 All Halodule along the North American coast have been considered to be H. beaudettei (C. den Hartog 1964, 1970; concepts accepted by D. S. Correll and H. B. Correll 1972 and R. K. Godfrey and J. W. Wooten 1979). A study of the morphology of Halodule in one large population in the northern Gulf of Mexico showed the leaf tips in one population to range from that of H. beaudettei to that of H. wrightii (R. C. Phillips 1967). A later study of the isozymes of the two morphologic types showed no difference between the two (C. McMillan 1991). I amWe are following both R. C. Phillips and C. McMillan in accepting one species of Halodule in the Gulf of Mexico, i.e., H. wrightii. Halodule beaudettei, therefore, is a synonym of H. wrightii.

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Names and Taxonomy

Taxonomy

Comments: Kartesz (1994) includes Halodule wrightii in H. beaudettei but the former name has precedence over H. beaudettei (FNA 2000).

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