Ascidians of South Padre Island, Texas, with a key to species.Abstract. -- The ascidians of South Padre Island, Texas South Padre Island is a resort community in Cameron County, Texas, United States [1] As of the 2000 census, the town had an official population of 2,422, although this significantly underestimates its size since many people split their time between vacation properties were surveyed in August 2004. Because the subtidal area is limited to soft sediments, the survey was restricted to marina floats and pilings, harbor buoys, boat hulls and other artificial substrates which offer suitable attachment surfaces for ascidians. Fifteen species were documented, with multiple species representing each of the three orders of ascidians. None of the species found in this survey are native, suggesting they were all introduced through boat traffic. About half the species were found in a reproductive state, however, indicating that they have established local breeding populations. ********** Ascidians are marine invertebrate invertebrate (ĭn'vûr`təbrət, –brāt'), any animal lacking a backbone. The invertebrates include the tunicates and lancelets of phylum Chordata, as well as all animal phyla other than Chordata. chordates, some of which are classic model organisms for the study of development and evolution (Conklin 1905; Berrill 1932; Satoh 1994; Corbo et al. 2001). They are emerging model organisms for other fields, including genetics (Dehal et al. 2002; Satoh et al. 2003), immunology (Azumi et al. 2003; Khalturin et al. 2003; Du Pasquier 2004; Rinkevich 2004), and neurobiology Neurobiology Study of the development and function of the nervous system, with emphasis on how nerve cells generate and control behavior. The major goal of neurobiology is to explain at the molecular level how nerve cells differentiate and develop their (Meinertzhagen & Okamura 2001; Meinertzhagen et al. 2004). Ascidians are also attracting attention as potential bio-indicators of environmental health (Cima et al. 1995; Cima et al. 1997) and as seafood, particularly in Japan and Korea (Sawada et al. 2001). Ascidians are efficient filter feeders, and certain species with wide environmental tolerances have become highly invasive, especially in bays and harbors where they compete with and overgrow o·ver·grow v. o·ver·grew , o·ver·grown , o·ver·grow·ing, o·ver·grows v.tr. 1. To grow over with herbage or foliage. 2. To grow beyond or too large for. v.intr. commercial shellfish (Lesser et al. 1992; Carver et al. 2003) and create a significant fouling community on boat hulls and marina floats (Teo & Ryland 1995; Hodson et al. 2000; Lambert 2001; 2002; Lambert & Lambert 2003). Thus, locales with high ascidian populations hold great potential for scientific and commercial research. Most ascidian species require a hard substrate for attachment. The natural subtidal substrates along most of the Texas coast are composed of soft sediments. Thus, prior to the establishment of man-made substrates (marina floats, pilings, harbor buoys and boat hulls), few shallow-water ascidians were recorded from the Texas Gulf coast (Van Name 1945; Whitten et al. 1950; Van Name 1954). Informal observations indicate that the south Texas coast may support ascidians in greater abundance than the rest of the Texas coastline. This paper lists the 15 species observed during a recent survey around South Padre Island, their locations and abundance, and includes a taxonomic key to species. METHODS Individuals were collected from the waters of the Laguna Madre around the southern end of South Padre Island, Texas, on 7-8 August 2004. Collection locations were identified using the global positioning system Global Positioning System: see navigation satellite. Global Positioning System (GPS) Precise satellite-based navigation and location system originally developed for U.S. military use. (GPS). Figure 1 shows the six collection sites: (a) Sea Ranch marina (26[degrees] 4' 33.4" N, 97[degrees] 9' 52.8" W); (b) Parrot Eyes marina (26[degrees] 8' 0.4" N, 97[degrees] 10' 36.9" W); (c) Laguna Madre boat canal mid-channel buoy (26[degrees] 4' 1.2" N, 97[degrees] 10' 0.6" W); (d) the Coastal Studies Lab seawater intake support (26[degrees] 4' 4.9" N, 97[degrees] 9' 49.1" W); (e) Port Isabel deep water docks (26[degrees] 3' 30.0" N, 97[degrees] 12' 49.4" W), and; (f) Billy Kenan's dock (26[degrees] 3' 56.8" N, 97[degrees] 12' 54.6" W). Specimens were initially examined live under dissecting microscopes, with further examination of some species after preservation. Representative individuals were fixed either directly in 70% ethanol or relaxed in seawater containing a few drops of a concentrated menthol/ethanol solution, and then preserved in 10% seawater formalin formalin /for·ma·lin/ (for´mah-lin) formaldehyde solution. for·ma·lin n. An aqueous solution of formaldehyde that is 37 percent by weight. buffered with sodium borate sodium borate n. A crystalline compound that is the sodium salt of boric acid and is used as an alkalizing agent and as a mild astringent in lotions, gargles, and mouthwashes. . [FIGURE 1 OMITTED] Specimens were identified at least to genus level. The primary sources used for identification were (Van Name 1945; Plough 1978); Didemnum duplicatum was identified from (Monniot 1983). Labelled voucher specimens were deposited in the Coastal Studies Laboratory on South Padre Island. RESULTS Fifteen species of ascidians were identified in this survey (Table 1). Styela plicata, S. canopus, and Lissoclinum fragile were particularly abundant, with S. plicata being found in large numbers at four of the six collection sites. Several other species that are small or inconspicuous may also be more common or abundant than indicated by this survey. Seven species were reproductive at the time of the survey, indicating that these species have formed locally reproductive populations. Ascidia interrupta, though rare during this survey, is abundant in autumn and is reproductive during that time. KEY TO SPECIES "There are some groups of animals for which keys can be made that really work in a considerable number of instances, but the ascidians are not among them" (Van Name 1945). This key is specific for the organisms seen or previously collected in these bays but is not necessarily valid for other regions. It is based on a 7-9 August 2004 survey of South Padre Island; there may be additional species more abundant at other times of the year that are not included here. An asterisk (*) indicates species not found during this survey but which are expected due to their distribution: Ciona intestinalis has a cosmopolitan distribution, and Molgula manhattensis has been recorded elsewhere in Texas. Explanations of terms, species descriptions, and illustrations can be found in Van Name (1945) or Plough (1978).
1. Solitary ascidians; each zooid enclosed in 2
its own tunic
Colonial ascidians; multiple zooids within a 6
common tunic or connected by stolons
2. Branchial sac without internal longitudinal folds 3
Branchial sac with four or more prominent 4
internal longitudinal folds
3. Body wall (easily visible inside smooth Ciona
transparent tunic) with five to seven white intestinalis*
wide longitudinal muscle bands on each side
(often somewhat contracted in fixed animals);
animal elongate, flaccid, attached basally
Body wall muscles in a meshlike pattern mostly on Ascidia
right (uppermost) side but not as above; animal interrupta
attached broadly on left side, tunic
semi-transparent, thin and not smooth
4. Tunic thin, semi-transparent but usually muddy; Molgula
body spherical, 2-4 cm in diameter; oral siphon manhattensis*
with six lobes, atrial siphon with four lobes;
six branchial folds per side
Tunic leathery; four branchial folds per side 5
5. Tunic brownish, furrowed; body usually 2-3 cm in Styela canopus
height; siphon tips with numerous mottled
reddish stripes; two long slender ovaries/side;
testes large, white, often bifurcated, attached
to posterior end of ovaries by long threadlike
sperm ducts
Tunic white with large rounded soft lumps; body Styela plicata
up to 10 cm in height; siphon tips with four
black stripes; two gonads on left side, five on
right; testes small and attached along most of
the length of each ovary
6. Multiple zooids connected by stolons, each zooid 7
enclosed by separate tunic
Multiple zooids all embedded in common tunic 9
7. Zooids spherical or up to twice as long as wide 8
Zooids over four times as long as wide, Clavelina oblonga
transparent, colorless.
8. Tunic soft and fragile, zooids globular, pale Perophora sp.
green, translucent, 2-4mm in height; branchial (probably P.
sac with four rows of stigmata viridis)
Tunic tough and leathery, zooids elongate, dark Polyandrocarpa
brown or purple, up to 2 cm in height; stolons zorritensis
usually coalesced into a basal mat; branchial
sac with more than four rows of stigmata
9. Zooids not divided into body regions; vascular 10
ampullae present in tunic
Zooids divided into two or three distinct 13
regions; vascular ampullae absent in tunic
10. Zooids (2.5-4 mm) flat, never organized in 11
systems, widely spaced with clear tunic
between, both siphons open at colony surface
Small zooids (<2 mm), organized in systems, only 12
branchial siphon opens to surface of colony,
densely spaced with little tunic between
11. Zooids red, tunic opaque Symplegma rubra
Zooids translucent with greenish or multicolored Symplegma viride
flecks of pigment
12. Zooids in elongate systems, colony a single Botrylloides
color, usually purple or orange, vertically nigrum
oriented in tunic, testis ventral (on side with
incurrent siphon) and anterior to single ovary,
stomach lobes bulbous at ends
Zooids in elongate systems, two colors in colony, Botrylloides sp.
dark basic colony color, bright yellow around
siphonal area
13. Zooids with two body regions (thorax, abdomen), 14
colony thin and encrusting, zooids with four
rows of stigmata
Zooids with three body regions (thorax, abdomen, Polyclinum
post-abdomen), colony dark, thick and constellatum
encrusting, may be dome shaped, zooids in
circular systems, each zooid with 14-18 rows of
stigmata
14. Tunic with tiny (visible with compound 15
microscope) white spherical calcareous spicules
with many short pointed rays, mostly in surface
layer of colony
Colony lacking calcareous spicules though there Diplosoma
may be considerable white pigment granules; listerianum
tunic transparent, very flaccid, zooids tiny
(2-3 mm in length) usually with black pigment
on thorax and abdomen
15. Atrial opening small or moderate size; sperm 16
duct spirally coiled, colony not white and
easily torn
Atrial opening large, exposing most of branchial Lissoclinum
walls; sperm duct not spirally coiled, colony fragile
white, tunic very fragile and easily torn
16. Colony distinctly muddy gray colored due to Didemnum
numerous fecal pellets stored in the tunic psammathodes
Colony salmon colored, leathery, with meandering Didemnum
dark lines. duplicatum
DISCUSSION A diverse assemblage of ascidian species is present in considerable abundance along the southern Texas coastline. All of the species found in this survey are apparently non-native and have most likely been introduced on boat hulls. All have been recorded elsewhere in the Gulf of Mexico Noun 1. Gulf of Mexico - an arm of the Atlantic to the south of the United States and to the east of Mexico Golfo de Mexico Atlantic, Atlantic Ocean - the 2nd largest ocean; separates North and South America on the west from Europe and Africa on the east , on the Atlantic side of Florida, or various regions of the Caribbean as well as other warm water regions of the world (Lambert 2001; 2002). All are shallow-water species not recorded in the survey of (presumably pre·sum·a·ble adj. That can be presumed or taken for granted; reasonable as a supposition: presumable causes of the disaster. native) deep-water ascidians of the Gulf of Mexico (Monniot & Monniot 1987), though a few were recorded from continental shelf depths of the Gulf (Plough 1978). Given that five colonial species contained brooded larvae Larvae, in Roman religion Larvae: see lemures. , and two of the three solitary species had ripe gonads, it seems likely that many or most of the species found have formed breeding populations in the local waters. The species sampled include more than one member of each of the three orders in class Ascidiacea, providing substantial diversity for comparative research. Indeed, the prospects for future research on ascidians in this area are extremely good. Many of the genera found on South Padre Island have been the focus of substantial research. For example, the natural pigmentation pigmentation, name for the coloring matter found in certain plant and animal cells and for the color produced thereby. Pigmentation occurs in nearly all living organisms. of Styela embryos enabled classic studies of chordate chordate Any member of the phylum Chordata, which includes the most highly evolved animals, the vertebrates, as well as the marine invertebrate cephalochordates (see amphioxus) and tunicates. development (Conklin 1905; Gehring 2004). Colonial tunicates like Botrylloides are now model organisms for allorecognition and the evolution of immune responses (Scofield et al. 1982; Scofield & Nagashima 1983; Rinkevich 1995; Hirose et al. 1997; Paz & Rinkevich 2002; Rinkevich 2004). Several of the ascidian genera on South Padre Island have been the source of many novel chemical compounds, including some with possible therapeutic properties, including Didemnum (Kang & Fenical 1997; Smith et al. 1997; Davis et al. 1999; Mitchell et al. 2000; Oku et al. 2003), Lissoclinum (Badre et al. 1994), Styela (Lee et al. 1997a; Lee et al. 1997b; Zhao et al. 1997) and Symplegma (Lindsay et al. 1999). The collecting sites are conveniently located near a well-equipped research and teaching laboratory (Coastal Studies Laboratory, University of Texas-Pan American). All the species described here should be easily maintained alive in the large seawater tanks, especially if placed in floating plastic sieves or grown on glass plates, or easily collected for same-day use. Most of the species have long breeding seasons and are easy to remove gametes from (for solitary species) or brooded embryos (for colonial species). 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FHL was founded in 1903 by University of Washington Zoology Professor Trevor Kincaid. 620 University Road, Friday Harbor, Washington Friday Harbor is a town in San Juan County, Washington, United States. The population was 1,989 at the 2000 census. Located on San Juan Island, it is the major commercial center of the San Juan Islands archipelago. 98250, Department of Biology, University of Texas-Pan American 1201 W. University Drive, Edinburg, Texas 78541 and Department of Carcinogenesis car·ci·no·gen·e·sis n. The production of cancer. carcinogenesis production of cancer. biological carcinogenesis viruses and some parasites are capable of initiating neoplasia. , University of Texas M.D. Anderson Cancer Center, Smithville, Texas 78957
Table 1. Systematic listing of species collected. Locations as given in
Methods and Figure 1.
Location(s) Solitary or
Taxon collected Colonial
Phylum Chordata
Subphylum Tunicata
Class Ascidiacea
Order Aplousobranchia
Family Didemnidae
Didemnum psammathodes b Colonial
Didemnum duplicatum b, c Colonial
Diplosoma listerianum b Colonial
Lissoclinum fragile a, b Colonial
Family Polyclinidae
Polyclinum constellatum b, e Colonial
Family Clavelinidae
Clavelina oblonga d Colonial
Order Phlebobranchia
Family Perophoridae
Perophora sp. a, e Colonial
Family Ascidiidae
Ascidia interrupta a Solitary
Order Stolidobranchia
Family Styelidae
Botrylloides nigrum b, e Colonial
Botrylloides sp. c, e, f Colonial
Polyandrocarpa e, f Colonial
zorritensis
Styela canopus a, b, e Solitary
Styela plicata a, b, e, f Solitary
Symplegma viride e Colonial
Symplegma rubra a, e, f Colonial
Reproductive Statis
Taxon during Survey
Phylum Chordata
Subphylum Tunicata
Class Ascidiacea
Order Aplousobranchia
Family Didemnidae
Didemnum psammathodes Not productive
Didemnum duplicatum Brooded larvae
Diplosoma listerianum Brooded larvae
Lissoclinum fragile Not productive
Family Polyclinidae
Polyclinum constellatum Not productive
Family Clavelinidae
Clavelina oblonga Brooded larvae
Order Phlebobranchia
Family Perophoridae
Perophora sp. Not productive
Family Ascidiidae
Ascidia interrupta Not productive
Order Stolidobranchia
Family Styelidae
Botrylloides nigrum Not productive
Botrylloides sp. Brooded larvae
Polyandrocarpa Not productive
zorritensis
Styela canopus Ripe gonads
Styela plicata Ripe gonads
Symplegma viride Not productive
Symplegma rubra Brooded larvae
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