Associated with Marine Vertebrates in Puerto Rico and Florida

by

Jamilette Cintrón-De Jesús

A thesis submitted in partial fulfillment of the requirements for the degree of

Master of Marine Science

In

Biological Oceanography

University of Puerto Rico Mayagüez Campus 2001

Approved by:

Ernest H. Williams, PhD Date President, Graduate Committee

Antonio A. Mignucci- Giannoni, PhD Date Member, Graduate Committee

Juan G. González Lagoa, PhD Date Member, Graduate Committee

Dallas E. Alston, PhD Date Director of the Department of Marine Science

Angel Berrios, PhD Date Representative of Graduate School

Jaime Seguel, PhD Date

1 2 Introduction

Most studies about cirripeds or barnacles are directed toward finding methods for their removal from marine structures and vessels. Other studies focus on biodiversity, phylogeny, zoogeography and life history parameters of .

Barnacles (balano, ballocas or caracolillos, in Spanish) are belonging to the subclass Cirripedia and to the order

(true barnacles). They can be divided in two groups by their types and morphological differences (Fig. 1): The acorn and stalk or “goose” barnacles. Some species are edible and commercially managed (percebes, in Spanish). They are mostly marine but some species are resistant to fresh water, i.e., improvisus (Zullo, 1979).

Many species are located in the intertidal zone and can usually be observed in piers, seawalls, marinas and boats. For that reason most of the studies have been directed to finding ways or eradicate them. In boats and ships, they increase fuel consumption. This research mostly looks at the adhering mechanics and morphology of the cirripeds. Barnacles are mostly hermaphrodites, but they need another individual to fertilize them. They are usually gathered in clusters to assure their reproduction. A single barnacle can produce thousands of eggs (Poor and Taylor, 1998). Eggs hatch into microscopic nauplii larvae that pass through six metamorphic changes each increased complexity until the cyprid crawls over a surface with the help of its 3 antennae and attaches from the head with a special cement or glue excreted by the antennae. At that point, the cyprid stage changes to the juvenile, begins to develop and turns in to a sessile crustacean. If larvae do not find a place to settle in a limited amount of time, they cannot complete their cycle and will die.

Some species from the order of Rhizocephala are parasitic. The rest have a symbiotic relationship known as commensalism. Barnacles feed by sweeping the water with their cirri, or feathery legs. The mechanics of this movement depends on the species. Some maintain their operculum (the opening from where the cirri protrude) closed tightly and others partially closed. They can extend their cirri two or three times per day or only in high tides or only during the nights, depending on the species. The species, and sometimes the environment, determines essentially the feeding mechanisms of the barnacle.

The literature concerning the of barnacles in the Caribbean, and thus Puerto Rico, is scarce. Mignucci-Giannoni (1996) reported these to be epibionts of marine vertebrates, specifically cetaceans, sirenians and chelonids, confirming their existence, but not establishing a detailed list of their taxonomy identification, characteristics or description.

It has been presumed that some barnacle species are specific to a certain substrate or host. This research examines the correctness of these assumptions, providing a description of each species and a key of the 4 cirripeds found associated with the West Indian (Trichechus manatus), the (Megaptera novaeangliae), the green sea (Chelonia mydas), and the hawksbill sea turtle (Eretmochelys imbricata). Additionally the samples were examined to verify occurrence, geographic range and morphometric comparisons of the shell diameter and height and operculum measurements of each barnacle.

Literature review

Naturalist Sir Robert Morey (1678) from Great Britain referred to the cirripeds as “little birds perfectly shaped” believing that barnacles were eggs of barnacle geese. When Linnaeus (1758) began to classify the kingdom, he considered barnacles to be mollusks. Later, Vaughn Thompson, a British army surgeon, demonstrated that barnacles were crustaceans.

Charles Darwin studied in his two monographs, cirripeds and wrote a monograph on these, divided in two parts, or pendunculate cirripeds (Darwin 1851), and Monograph of the , the Verrucidae

(Darwin 1854). These are still the most important reports on barnacles. Many cirripedologists still use these references for the classification and description of the species.

Many others studies have been conducted concerning the biology of barnacles, but only in the last decade the physiology and the ecology of barnacles has begun to be studied. Most works are about removal of 5 barnacles from ships due to economics limitations that these cause on fuel efficiency. Others contribute with geographic location and classification.

Pilsbry (1916) classified the barnacles in the U.S. National Museum collections, while Nilsson-Cantell (1927) classified barnacles in the British

Museum.

Parker (1960) studied the ecology of cirripeds, and detailed characteristics of some barnacles. Crisp (1967) studied how substrate influenced barnacle morphological development, as well as details on phylogeny of the group. Newman (1976) revised the taxonomy of balanomorph barnacles, including a catalog of the species. Zullo (1979) reported on the marine flora and fauna of the northeastern United States, including the characteristics of barnacles in New England. The same year,

Monroe and Limpus (1979) provided the systematics and the characteristics of all barnacle species found on in Queensland, Australia. Anderson

(1994) provided an extensive list of references and excellent information of the biology and phylogeny of the cirripeds.

In the Gulf of Mexico, Pilsbry (1953) detailed Florida barnacles, while it was not until the 1980s that Gittings (1986) studied barnacles in its northern part. In the Caribbean, especially in Puerto Rico, the knowledge about barnacles associated with marine vertebrates is minimal. While poorly,

Bigelow (1899) detailed the cirripedia collected by the Fishhawk Expedition in

1898-1899. Williams (1978) and Williams et al. (1986) reported on the 6 associations of pendunculate barnacles with fishes, one from the Japan area and the other in the Caribbean area off St. Thomas. Rodríguez-Fourquet

(1992) studied the barnacles present in the estuarine ecosystems of Joyuda

Bay.

In the 1990s, the Caribbean Stranding Network began their operations, and as a consequence, major studies concerning marine mammals and sea turtles increased. As part of their mortality assessment research, barnacles were collected from stranded and stored for future analysis. Mignucci-Giannoni (1996), Mignucci-Giannoni et al. (1998),

Mignucci-Giannoni et al. (1999), Rodriguez-López and Mignucci-Giannoni

(1999), and Rosario-Delestre et al. (1999) reported the presence of barnacles on marine mammals from Puerto Rico and the Virgin Islands.

Conchoderma auritum, sp. and manati were identified from the samples. Materials and Methods

Barnacles were collected and removed from stranded whales

(humpback whale, Megaptera novaeangliae), (Trichechus manatus) and sea turtles (hawksbill sea turtle, Eretmochelys imbricata, and , Chelonia mydas) in Puerto Rico and Florida. They were collected during necropsy or salvage procedure following the protocols of

Mignucci-Giannoni et al. (1998) and Ortíz-Rivera (2000).

Most samples were preserved in 70% ethanol, except for those found in humpback whales, which were fixed first in 10% formalin and later preserved in ethanol. Barnacles examined for size and strength of the shell were cut with a geologic saw. Remaining samples from manatees and sea turtles were dissected using pliers. A small brush was used to clean the barnacles from inorganic and organic material.

Identification and classification of the barnacles was done using

Darwin (1854), Pilsbry (1916), Newman and Ross (1976), Gittings et al.

(1986) and Anderson (1997) as reference. Each barnacle specimen was identified based on morphological differences, primarily external characters.

Some were dissected and observed internally with a stereoscope.

Specimens were measured with calipers to determine their shell diameter, shell height and diameter of the operculum. Operculum/shell diameter and

7 8 shell height/shell diameter were used for statistical analysis and to provide a better morphological description.

All data were analyzed using t-test (two populations) independently.

The comparison was done between locations with same host and different host's species in the same location. Box chart graphs were used, where each

Y column of data is represented as a separate box. The column names or labels supply the X-axis tick labels. The horizontal lines in the boxes denote the 25th, 50th and 75th percentile values. The error bars denote the 5th and

95th percentiles values. The two symbols below the 95th percentile error bar denote the 0 and 1st percentile values. The two symbols above the 95th percentile error bar denote the 99th and 100th percentiles. The square symbol in the boxes denote the mean of the column of data.

Results

A total of three barnacles from humpback whales, (Megaptera novaeangliae) one from Florida and two from Puerto Rico, were examined.

Two barnacle species were found directly attached to the skin of the whale.

These were and Coronula reginae (Table 1). A third one, a pendunculate barnacle, Chonchoderma auritum was found attached only to

Coronula diadema (Table 1). Coronula diadema was the most common 9 barnacle in this whale in comparison with the four individuals of Coronula reginae found. (Table 2). Both species occurred on the same whale.

A total number of six sea turtles were examined. from Puerto Rico.

The sea turtles examined were the green turtle, (Chelonia mydas) and the hawksbill turtle, (Eretmochelys imbricata) (Tables 1, 3). The barnacles species on the sea turtles were, Stomatolepas dermochelys and hexastylos; and Chelonibia caretta was on the hawksbill, (Table 1, 3). All of them can coexist in the same host but it is more common to find P. hexastylos.

Manatees Trichechus( manatus) included 45 individuals, 6 from

Puerto Rico and the rest from Florida (Table 4). Barnacles species found attached directly to the skin of the manatee were, P. hexastylos , P. decorata and a modified Chelonibia manati subspecies and Chelonibia patula (Table

1, 4). Balanus eburneus and B. amphitrite (Table 1, 4) were found over the

C. manati not directly on the skin of the manatee. C. manati lobatosis or crenatibasis

(Table 1, 4) (Pilsbry, 1916) occurred more commonly in the Florida manatees than the Platylepas species .

A brief summary of the barnacle species morphometrics per host can be found in Tables 5 to 7. Table 5 compares the difference between both

Coronula and compares the same species on different localities. Table 6 compares barnacle species over two different sea turtle species from Puerto 10 Rico and Table 7 show the morphometrics differences on barnacles from different locations.

Barnacle Descriptions

Coronula diadema (Linnaeus)

“The most common barnacle found on humpback whales in the

Atlantic”.

(Fig. 2)

Name – Linnaeus (1767) was the first referring to this barnacle as

diadema. De Blainville (1824) was the first that use the ,

Coronula.

Diagnostic characters – Shell forming a crown and tall. Operculum larger

than the base and hexagonal. Radii broad an tall, edges crenated,

terga absent or vestigial. The underside of the shell is deeply

concave (Darwin). Ribs broad and rounded. Diameter from 44 until

86 mm and height from 30 to 60 mm.

Records – One hundred thirty-seven specimens of this species were found

on a total of 141 individuals. One hundred thirty-five (135) were

found in Puerto Rico and the rest in Florida.

Geographic range – Found on whales from the east of United States, in the

Arctic Seas, the Atlantic Ocean and the Gulf-Stream. According to

Newman and Ross (1976) it is cosmopolitan. 11 Ecology – From oceanic waters.

Associations – Commonly found associated with auritum.

Also small crustaceans from the subclass Branchiura can be found

over and inside the barnacles from the Rincón and Naguabo whales.

Location on the host – Majority located near the jaw, mouth, all over the

face and flippers, but can be found sporadically in the rest of the

body.

Average diameter and height measured on the samples– Florida, 42.75

mm-22.65 mm. Puerto Rico, 40.27 mm-23.41 mm.

Host Specificity – Found only on whales, but not specifically on the

humpback whale. Some records from gray whale and others

cetaceans (Newman and Ross, 1976).

Coronula reginae (Darwin)

Some taxonomists believe that Coronula reginea is the most

common barnacle found on whales in the Pacific Ocean, dominating

that ocean like

C. diadema dominates the Atlantic Ocean (Fig. 3).

Name-Reported and described by Darwin (1854).

Diagnostic characters-Shell rounded or depressed at the upper part. Height

lower than C. diadema. Longitudinal and much flattened ribs

(Darwin) than in C. diadema. Operculum hexagonal but apparently

a little smaller than C. diadema and the terga is absent like C. 12 diadema. Diameter from 48 to to 60 mm and height from 13 to 19

mm (Pilsbry, 1916).

Records – Found on one humpback whale on the coast of Rincón, PR.

Geographic range-Found mainly on whales in the Pacific Ocean and according to Pilsbry (1916) in some whales of the Northern Atlantic.

Ecology-Oceanic waters.

Associations –No associations reported.

Location on the host-– Majority near the jaw, mouth, all over the face and

flippers, but can be found sporadically in the rest of the body.

Average diameter and height measured on the samples – 35.55 mm-

14.73mm.

Host Specificity – Apparently on all cetaceans, from the Pacific.

Chelonibia caretta (Spengler)

Generally attached to the carapaces of sea turtles. (Fig. 4)

NAME – WAS IDENTIFIED BY SPENGLER AS LEPAS CARETTA AND

DARWIN ADDED THE GENUS CHELONIBIA IN HIS MONOGRAPH.

Diagnostic characters-Shell extremely strong, heavy, broadly oval, white.

Operculum (orifice) oval, not at all angular. (Darwin, 1854) The most

diagnostic character is that the radii is absent or very narrow

presenting only lines of suture. Their compartments have simetry. 13 Their basal edges finely dentate. The rostrum and the carina not

facing each other. Greatest diameter 22 mm and height of 8 mm.

Records – Only three specimens were found on a total of forty-five turtles.

All from a hawksbill sea turtle (E. imbricata).

Geographic Range – Tropical and subtropical species. Reported from

Venezuela, Africa, Northern Australia, Brazil and the West Indies.

Ecology – Difficult to establish but from oceanic or coastal waters.

Associations-Apparently no associations with other organism occurs.

Location on the host – Over the carapace of the sea turtle, especially the

front area.

Average diameter an height measured– 28.07 mm and 7.53 mm, samples

only from Puerto Rico.

Host Specificity – Apparently on all the sea-turtles. Found exclusively on

samples of E .imbricata.

Chelonibia patula (Ranzani)

Name-Described by Ranzani (1918). Later Pilsbry (1916) used the same

name in his descriptions. (Fig. 5)

Diagnostic characters-Shell light brown or white, fragile, the radii thin.

Diameter of the operculum usually large, basal margin of the shell

smooth, not embedded in the skin. The opercular valves narrower

than in other species. Greatest diameter 22mm and height 8mm. 14 Records-Found only in two manatees from Florida. A total of 31 barnacle

individuals.

Geographic range-Worldwide distribution in warm seas. (Zullo, 1979)

Ecology – Coastal and estuarine waters.

Associations-no association was found with other animals, most common

host are .

Location on the host-Not reported. Apparently on the back of the manatee.

Average diameter and and height measure-13 4 and 8.2mm

Host Specifity-Originally reported on crabs, later reported for sea turtles

(Pilsbry, 1916) . Found on manatees samples only. This is a new

host record.

Platylepas hexastylus (Fabricius)

Name – Fabricius first described this species in 1789. He named it Lepas

hexastylus. Later Darwin (1854) refer to the same species as

Platylepas bissexlobata. Pilsbry (1916) used

for the same species (Fig. 6).

Diagnostic characters-Shell dome-shaped, broadly oval or circular.

Operculum oval Tha basal membrane forms a convex layer. The

membranous basis being streched over them. Midribs not

prominent. Diameter between 9 to 10 mm in Florida (Pilsbry, 1916) 15 Records-Three were found on seven manatees in Puerto Rico and in one of

forty-four manatees in Florida. This species was also found on both

species of sea turtles, E. imbricata and C. mydas.

Geographic range-is found worldwide in tropical and warm temperate seas

on turtles, and manatees.

Ecology-Apparently from coastal calm waters.

Associations-P.hexastylus can coexist with other barnacles like C. caretta.

Apparently can coexist but in minor quantity with C. manati

lobatosis. Probably they are competitive for the same space in the

host. These were extraordinary conspicuous in Puerto Rican

samples.

Location on the host – All over the body of the host, but especially over

their back.

Average diameter and height measured– On manatees from Florida, 9.70

mm and 3.57 mm. On manatees from Puerto Rico, 3.89 mm and

height of 1.24 mm. On sea turtles in Puerto Rico, the

measurements were 4.13 mm for the E. imbricata and 3.93mm for

C. Mydas.

Host Specificity-One of the most widely distributed on marine vertebrates,

can be found on sea turtles, manatees and dugongs and sea

snakes (Pilsbry, 1916).

Platylepas decorata (Darwin) 16 Name – Same as P. multidecorata that Daniel (1962) described. Was

originally described by Darwin (1854) in his monograph of the

cirripeds (Fig. 7).

Diagnostic characters – Shell subcircular, with fine longitudinal ridges,

ornamented in the lower part with small beads, ring like, basal

membrane in convexity. Conspicuous midribs.

Records – Found on manatees in both geographic areas studied, but not in

the sea turtles although noted in them by Monroe and Limpus (1979)

Geographic range-From Galápagos, through Pacific Oceania to the western

coast of Australia, and in the Atlantic Coast of North America and

the Caribbean islands.

Ecology – Apparently in all ocean waters.

Associations – Can coexist on the host with the other, Platylepas

hexastylus and with the different barnacles of the genus of Balanus.

Location on the host – Anywhere, but especially over their back.

Average diameter and height measured – In Puerto Rico 5.65 and 2.61

mm. In Florida 8.42 and 2.85 mm.

Host Specificity – According to Monroe and Limpus (1979) this species was

found on sea turtles in their research but they noted that it can occur

on other hosts. Found on manatees in both geographic areas.

Stomatolepas dermochelys (Costa) 17 Found at least one specimen on both species of sea turtles. (Fig. 8)

Name – Costa described as Coronula elegans. It was reported as S.elegans

prior to renaming by Monroe and Limpus (1979).

Diagnostic characters-Bowl shaped shell, operculum more wide than the

base, no internal midribs. (Gittings et al., 1986). The shell has

scales composed of small oval convolutions. A triangular area bare

of scales at the middle bottom edge of each compartment forms a

chevron pattern (Monroe and Limpus, 1979) Scuta and terga From

6 to 9 mm diameter and height from 3 to 6 mm.

Records –Three specimes found on two different sea turtle species. One in

C. mydas and two in E. imbricata.

Geographic range – Worldwide in warm seas.

Ecology-From coastal waters to oceanic

Associations – Found together (coexist) with P. hexastylos on the same

host. No apparent association with other organims in the samples

obtained.

Location on the host – In the soft skin, mouth and throat of sea turtles.

Average diameter and height measured– In E. imbricata, 3.95 and

1.40mm. In C.mydas 3.00 and 1.30mm.

Host Specificity – Apparently specific to sea turtles. Was found on the sea

turtles samples of E.imbricata and C. mydas.

Balanus eburneus-(Gould) 18 Name-Reported by Gould (1841) on the Invertebrata of Massachussets.

(Fig. 9)

Diagnostic characters – White to yellow shell. Obliquely conic or shortly

cilindric (Pilsbry, 1916), smooth with the operculum acute. Covered

with yellow epidermis, the radii white without epidermis. (Darwin,

1854). Scutum striated longitudinally. The tergum is broad, without

external furrow. The carinal –margin protuberant in the superior part,

highly arched. Their radii and alae are wide, with oblique summits.

Records-Reported on 10 manatees from the area of Florida only.

Geographic range – Endemic to western Atlantic, from Boston until Rio de

Janeiro; introduced to Europe, Mediterranean, Indian ocean, Pacific

Ocean, Panama, Jamaica and West Indies.

Ecology – Most abundant in estuaries and nearshore.

Associations-Apparently needs a hard substrate to attach. Uses other acorn

barnacles like C.manati for that purpose.

Location on the host-Do not attach directly to the manatee. Use the

substrate of other barnacles and those are their real hosts. When the

population is too high it sometimes can be found attached partially of

some barnacle and to the skin of the manatee.

Average diameter and height measured – 4.95 and 4.10 mm in Florida

only. 19 Host Specificity – Found only directly associated with the C. manati .

Probably can be found over other Chelonibia species that attach to

other hosts.

Balanus amphitrite-(Darwin)

Widely distributed Balanus, numerous subspecies (Fig. 10).

Name – Reported first by Wood as Lepas radiata in 1815. In 1826 Risso

reported the same species as Balanus balanoides. Finally Darwin

(1854) used Balanus amphitrite in his classification.

Diagnostic characters – Shell longitudinally striped with purple or pink,

sometimes with the stripes confluent. Sometimes completely white.

Records – Only one specimen was identified on a manatee from Florida.

Geographic range – This species is considered cosmopolitan in tropical and

warm seas.

Ecology – Common in coastal waters.

Associations-. Commonly associated with the other barnacle B.

tintinnabulum (Darwin)

Location on the host-Not directly attached to marine vertebrates.

Diameter and height measured – 6.00 and 3.40 mm

Host Specificity – Not specific. They are extremely common on ships

bottoms and often over floating items like buoys.

Chelonibia manati (Gruvel)

C. m. lobatosis or C. m. crenatibasis (Pilsbry) 20 Name – first identified in 1903 by Gruvel. Pilsbry (1916) added in his

cirripeds classification, two “new variety” subspecies. (Fig. 11)

Diagnostics characters – (Pilsbry, 1916) For Chelonibia manati – sides of

the well develop radii not dentate, the parietes having salient

longitudinal ribs which subdivide toward the base, and the basal septa

are fewer.

For C. m. lobatosis – has a large operculum and steep walls, which are

irregularly plicate or ribbed in the lower half, producing an irregularly

lobed periphery. The radii are rather wide, not dentated. Diameter 23

mm and height 9.5 mm.

For C. m. crenatibasis – has a very large operculum, their radii partly

simple, but a few show very weak traces of “teeth”. Very close to the

base there are numerous inconspicuous folds and the interstices form

small loops projecting into the septate base. Diameter from 43 to 37

mm and a height from 18 to 13 mm.

Records – Found in three of six manatees in Puerto Rico and in all

manatees of Florida.

Geographic Range – (Newman and Ross)

Chelonibia manati – West Africa

C. m. lobatosis – Florida

C. m. crenatibasis-Unknown 21 Ecology – Apparently from coastal and estuarine waters.

Associations – A large group of Balanus can use the Chelonibia manati

shells to establish themselves.

Location on the host – On manatee skin, all over the body but with mainly

on the back of the animal.

Average diameter and height – In Puerto Rico – 10.21 and 4.98 mm. In

Florida – 13.13 and 5.86 mm.

Host specificity – Pilsbry (1916), Newman and Ross (1976) found C. m.

lobatosis only on Caretta caretta (loggerhead). Here was found

exclusively over manates.

Discussion

Nine box chart graphics were done to compare the populations between hosts on different geographical locations. P. hexastylos (Figs. 12,

13) on different sea turtles demonstrated that they are not significantly different. The same occurred with P. decorata (Figs. 17, 18). However for this species the operculum with respect of his diameter on the E. imbricata is lower compared with the C. mydas . With the height is inverse. The

Stomatolepas dermochelys species (Tables 3, 6) was not analyzed, their n is only three. On humpback whales the only species analyzed was C. diadema.(Fig. 14), results showed a pattern similar to sea turtles, no significant differences were found. 22 Species analyzed for manatee samples were P. hexastylos

(Figs. 19, 20). and the subspecies C. manati (lobatosis or crenatibasis) (Figs.

15, 16). P. hexastylos ratios never showed significant differences. C. manati lobatosis species present the characteristics that Pilsbry (1916) mentioned in his research for the same species except for one, that their radii is without

“teeth”, At the same time he mentioned the possibility of another subspecies named C. m. crenatibasis. whose base “ form minute loops and show very weak of traces of teeth ” in the radii. C. manati lobatosis showed characteristics of both subspecies.Observing our specimens of C. manati lobatosis, a possible pattern of stage development can be an additional information to understanding and classifing them. Differences in these shells are only changes in shape as they mature. (Fig. 11) . Statistical analysis of the population of barnacles from Florida and Puerto Rico showed a significant difference between these geographic areas. It is important to indicate that the occurrence of C.manati lobatosis is higher in Florida than

Puerto Rico. The “dominant” barnacle in manatees from Puerto Rico over is

P. hexastylos and in Florida is C.manati lobatosis or crenatibasis. This difference probably represents competition for space.

As shown in the Table 7 the species from Florida exhibit a diameter larger than the ones from Puerto Rico, but in contrast, the species from

Puerto Rico showed a greater height in comparison with the Florida ones. 23 Graphs show that these barnacles differ in each location. Probably C. m lobatosis and C. m. crenatibasis represent the same subspecies.

Differences with Florida can be attributeed for some external factors like pollution (Royo-Gelabert, 1994). Manatees from Florida are more in an estuarine environment and those in Puerto Rico in a saline environment, these may be enough difference to explain the differences in the barnacles.

Probably C. manati lobatosis is more tolerant to fresh water than probably

Platylepas.

C. patula (Tables 1, 4) occurred only on two manatees from Florida. This species has been reported from crabs and occasionally on sea turtles.

Trichechus manatus.is a new host for this barnacle. The barnacle on T. manatus differred from the descriptions of the barnacle in that the basal margins of C. patula were noted in the description to be “smooth” (Pilsbry,

1916). The barnacles on T. manatus had “ramifications” of their margins.

This variation may just be an adaptation to an usual host.

The most relevant findings were that C.patula was found on manatees, not just on crabs and sea turtles. (Williams and Porter) A new host reported. C. manati lobatosis and crenatibasis apparently are the same subspecies, but their stages development were never discussed. Based on simple observation of the shell parietes, radii and base, this species probably undergoes more variation during it’s adult development than any other barnacle found on this research. C. m. lobatosis is more dominant in Florida 24 than in Puerto Rico. On the other hand P. hexastylus took the same space of

C. m. lobatosis and C. m. lobatosis was more used by many Balanus spp. for anchoring than the Platylepas species. Gittings (1986) mentioned that probably C. m. lobatosis is endemic to Gulf of Mexico, however was found in the Caribbean.

Gittings found only S. dermochelys on loggerheads turtles (Caretta caretta). In my research, S dermochelys was found on different species of sea turtles (E.imbricata and C. mydas). Conclusions and Recommendations

Eleven barnacle species were found in all the samples analyzed. One was lepadomorpha (pendunculate) and the rest (acorn).

Three barnacle species were found on humpback whales. The same number was found on sea turtles, and six species were found in manatees. One barnacle species P. hexastylos was shared on different hosts (manatees and sea turtles).

The most relevant findings were that C.patula was found on manatees, not just on crabs and sea turtles. (Williams and Porter, 1964) A new host reported. C. manati lobatosis and crenatibasis apparently are the same subspecies, but their stages development were never discussed.

Based on simple observation of the shell parietes, radii and base, this species probably undergoes more variation during it’s adult development than any other barnacle found on this research. C. m. lobatosis is more dominant in Florida than in Puerto Rico. On the other hand P. hexastylus 25 took the same space of C. m. lobatosis and C. m. lobatosis was more used by many Balanus spp. for anchoring than the Platylepas species. Gittings

(1986) mentioned that probably C. m. lobatosis is endemic to Gulf of Mexico, however was found in the Caribbean.

Gittings found only S. dermochelys on loggerheads turtles (Caretta caretta). In my research, S dermochelys was found on different species of sea turtles (E.imbricata and C. mydas).

The literature and the records in Puerto Rico about cirripeds is limited.

My research was basically about morphology differences and other data was sometimes incomplete. Data from the anatomical part of the host was lost when the barnacles were removed and some locations were not recorded.

Some data were obtained from sea turtles of different species, E. imbricata and C. mydas from Puerto Rico, but was impossible to compare them with

Florida samples because they were not collected. Using some knowledge about barnacles and the records in Florida, some comparisons were done it with the manatees. The statistical analysis showed that with the exception of

C. manati lobatosis, the barnacles in Florida and Puerto Rico are the same.

There is much additional research to be done in this geographic area.

Molecular, genetic and physiology research can be useful for classify and recognize some variation between species, and determinate if some new variety occurs in Puerto Rico or Florida area. 26 These crustaceans are interesting but they have been underestimated in their abundance and importance. Knowing where barnacles are more predominant, could possibly determinates migration routes of some of the marine vertebrates to which they commonly attach.

Analyzing the presence of heavy metals on the shells or tissues of the barnacles (Killingley and Lutcavage, ?) may indicate if the host was subject to pollution and other chemicals indicators of the health and exposure of their host. Direct or indirect more about these animals can contribute directly or indirectly to the information obtained from their hosts. 27

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Table 1. Barnacle species found in marine mammal hosts from Puerto Rico and Florida.

Host Barnacle species

Humpback whale (Megaptera novaeangliae) Coronula diadema Coronula reginae *

West Indian manatee (Trichechus manatus) Platylepas hexastylos Platylepas decorata Balanus eburneus* Balanus amphritrite* Chelonibia patula Chelonibia manati**

Hawksbill sea turtle (Eretmochelys imbricata) Chelonibia caretta Platylepas hexastylos Stomatolepas elegans

Green sea turtle (Chelonia mydas) Platylepas hexastylos Stomatolepas elegans

* species not directly attached to the host ** C. manati lobatosis or C. manati crenatibasis 33

Table 2. Barnacles species and number of individuals present on humpback whales (Megaptera novaeangliae) from Puerto Rico and Florida.

Locality Field no. Barnacle species No. individuals

Puerto Rico NEPST497 Coronula diadema 64 Coronula reginae 4 Puerto Rico NEPST509 Coronula diadema 70 Puerto Rico NEPST026 Coronula diadema 1 Florida Coronula diadema 2 Florida Coronula diadema 4

Table 3. Barnacles species and number of individuals present on sea turtles from Puerto Rico.

Field no. Barnacle species No. individuals

Hawksbill sea turtle (Eretmochelys imbricata) No number Chelonibia caretta 2 Platylepas hexastylos 1 NEPCH230 Chelonibia caretta 2 Platylepas hexastylos 6 NEPCH098 Platylepas hexastylos 1 NEPCH210 Platylepas hexastylos 6 Stomatolepas dermochelys 2

Green sea turtle (Chelonia mydas) NEPCH231 Platylepas hexastylos 16 NEPCH228 Platylepas hexastylos 10 Stomatolepas dermochelys 1

Table 4. Barnacles species and number of individuals present on West Indian manatee (Trichechus manatus) from Puerto Rico and Florida.

Locality Field no. Barnacle species No. individuals

Puerto Rico CPR9801 Platylepas hexastylos 10 Platylepas decorata 13 Puerto Rico CPR9702 Platylepas hexastylos 5 Puerto Rico TPR12 Platylepas hexastylos 4 Chelonibia manati * 8 Puerto Rico TPR11 Chelonibia manati* 11 34

Puerto Rico NEPST077 Platylepas decorata 4 Chelonibia manati* 2 Balanus eburneus 7 Balanus amphitrite 4 Puerto Rico NEPST532 Chelonibia manati* 12 Florida M81-11 Platylepas hexastylos 11 Balanus eburneus 7 Chelonibia manati* 39 Florida M81-50 Platylepas decorata 25 Chelonibia manati* 1 Florida M82-26 Balanus eburneus 4 Chelonibia manati* 6 Florida M81-28 Chelonibia manati* 70 Balanus eburneus 1 Florida M82-18 Balanus eburneus 1 Chelonibia manati* 19 Florida M81-67 Balanus eburneus 4 Florida M82-2 Balanus eburneus 4 Chelonibia manati* 16 Florida M81-29 Balanus eburneus 1 Chelonibia manati* 17 Florida M82-28 Balanus eburneus 1 Chelonibia manati* 1 Florida M84-15 Balanus eburneus 2 Chelonibia patula 4 Florida M81-26 Balanus eburneus 1 Chelonibia manati* 10 Florida M81-11 Balanus amphitrite 1 Florida M82-6 Chelonibia manati* 14 Florida M82-17 Chelonibia manati* 9 Florida M84-01 Chelonibia manati* 4 Florida M81-8 Chelonibia manati* 23 Florida M82-25 Chelonibia manati* 1 Florida M84-04 Chelonibia manati* 3 Florida M-82-3 Chelonibia manati* 6 Florida M81-23 Chelonibia manati* 23 Florida M84-28 Chelonibia manati* 12 Florida M82-30 Chelonibia manati* 6 Florida M86-09 Chelonibia manati* 7 Florida M82-24 Chelonibia manati* 4 Florida M82-12 Chelonibia manati* 8 Florida M82-19 Chelonibia manati* 10 Florida M82-11 Chelonibia manati* 9 Florida M79-26 Chelonibia manati* 5 Florida M81-58 Chelonibia manati* 19 Florida M82-16 Chelonibia manati* 9 Florida M86-8 Chelonibia manati* 8 Florida M81-15 Chelonibia manati* 14 Florida M81-14 Chelonibia manati* 48 Florida M81-12 Chelonibia manati* 10 Florida M82-22 Chelonibia manati* 8 Florida M81-69 Chelonibia manati* 21 35

Florida M82-32 Chelonibia manati* 11 Florida M80-21 Chelonibia manati* 66 Florida M81-65 Chelonibia patula 27

Table 5. Morphometrics of Coronula diadema and Coronula reginae on humpback whales (Megaptera novaeangliae) from Puerto Rico and Florida.

Puerto Rico Florida Morphometric Mean SD SEM Mean SD SEM

Coronula diadema Diameter (mm) 40.27 5.44 0.47 42.75 5.30 3.75 Opercular (mm) 23.41 3.44 0.30 22.65 1.20 12.02 Height (mm) 28.75 5.95 0.52 27.50 0.85 8.50

Coronula reginae Diameter (mm) 35.55 4.80 2.40 - - - Opercular (mm) 17.25 2.06 1.87 - - - Height (mm) 14.73 1.87 0.94 - - -

Table 6. Morphometrics of barnacles on different sea turtles from Puerto Rico.

Eretmochelys imbricata Chelonia mydas Morphometric Mean SD SEM Mean SD SEM

Platylepas hexastylos Diameter (mm) 4.13 1.11 0.30 3.93 0.91 0.18 Opercular (mm) 2.11 0.52 2.68 2.06 0.39 0.08 Height (mm) 2.80 2.68 0.77 2.53 2.68 0.53

Stomatolepas dermochelys Diameter (mm) 3.95 0.05 0.04 3.00 - - Opercular (mm) 1.80 0.10 0.07 1.50 - - Height (mm) 1.40 0.00 0.00 1.30 - -

Chelonibia caretta Diameter (mm) 28.07 9.l85 6.96 - - - Opercular (mm) 14.53 1.50 1.06 - - - Height (mm) 7.53 4.80 3.40 - - - 36

Table 7. Morphometrics of barnacles on West Indian manatee (Trichechus manatus) from Puerto Rico and Florida.

Puerto Rico Florida Morphometric Mean SD SEM Mean SD SEM

Platylepas decorata Diameter (mm) 5.65 1.77 0.44 8.42 1.74 0.35 Opercular (mm 2.61 0.57 0.14 3.67 0.56 0.11 Height (mm) 1.94 0.81 0.20 2.85 0.49 0.10

Platylepas hexastylos Diameter (mm) 3.89 1.63 0.37 9.70 1.31 0.40 Opercular (mm) 2.02 0.68 0.16 4.57 0.48 0.14 Height (mm) 1.24 0.44 0.10 3.57 0.46 0.14

Balanus amphitrite Diameter (mm) 10.10 7.64 1.37 6.00 - - Opercular (mm) 5.95 2.90 0.52 3.30 - - Height (mm) 4.50 4.95 0.89 3.40 - -

Balanus eburneus Diameter (mm) - - - 4.95 1.90 0.39 Opercular (mm) - - - 3.14 1.22 0.27 Height (mm) - - - 4.10 2.18 0.50

Chelonibia manati Diameter (mm) 10.21 6.83 1.21 13.13 5.81 1.10 Opercular (mm) 5.13 3.25 0.58 7.05 3.17 0.62 Height (mm) 4.98 3.44 0.62 5.86 3.30 0.69

Chelonibia patula Diameter (mm) - - - 13.40 6.13 1.10 Opercular (mm) - - - 8.20 3.48 0.62 Height (mm) - - - 8.27 3.83 0.69