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The effect of periodically "milking" to obtain Tyrian Purple from Plicopurpura pansa (Gould, 1853) on the frequency of expulsion and mortality.



ABSTRACT Most marine snails of the families Muricidae and Thaididae produce in their hypobranchial gland (mucus gland) a colorless secretion containing minute amounts of chromogens, which develop under the influence of light and oxygen into a pigment known as "Tyrian Purple." The hypobranchial gland of Plicopurpura pansa (Gould, 1853) is an exception among the muricids, because it is so active the snails can be stimulated periodically to expulse the secretion without harming the animals. In view of reported drastic declines of the populations of P. pansa because of the Tyrian Purple exploitation, in this laboratory study the effect of periodically "milking" of P. pansa on the frequency of expulsion and on the mortality of the snails was determined. At the beginning of the experiment using 110 animals (55 males and 55 females) only 30% expelled secretion. No relation was found between the occurrence of expulsion and the size or sex of the animals. In contrast to the laboratory snails the proportion of expulsions from free-living animals was 56%. Also here no differences were found between the occurrence of expulsion and the size or sex of the animals. For a period of 98 days it was tried in the laboratory to obtain daily secretions from 46 snails (23 males and 23 females). The frequency of expulsion declined drastically and at the end of the experiment no expulsions at all could be obtained. No difference could be noted between the sex and size of the snails and the decline of secretions, however, the total number of secretions during the test period was nearly double with the females, than with the males. The survival rate of the male snails was 83, of the females 87%. Handling mistakes during cleaning and feeding were the main reason for the mortalities. The attempt to milk the snails weekly caused, as during the daily milking experiment, a decline in the frequency of expulsion, and after 13 wk only 13% of the animals expulsed. To milk the animals every 2 wk caused a decline in the number of snails that expulsed, and only 18% of the animals secreted. No decline in the rate of expulsions was found by milking the animals every 3 and every 4 wk, and all animals survived. Frequent attempts to milk the animals has no impact on the survival rate, however, it affects the occurrence of expulsions. From the results reported here the decline of natural snail populations after milking could be explained by the environmental conditions of the intertidal zone The intertidal zone, also known as the littoral zone, in marine aquatic environments is the area of the foreshore and seabed that is exposed to the air at low tide and submerged at high tide, i.e., the area between tide marks. . After milking, the animals don't adhere fast enough to the rocks and they are washed away to unsuitable areas by the high wave action. For this reason, P. pansa should never be removed from the rocks, and the collection of the secretion should be allowed only in exceptional cases.

KEY WORDS: purple snail, Plicopurpura pansa, Tyrian purple

INTRODUCTION

The majority of purple-producing marine snails belong to the family of Muricidae, and most, if not all, produce in the hypobranchial gland (also called mucus gland) a colorless secretion, which is ejected into the mantle cavity, and turns on exposure to air and light to Tyrian purple (Fretter & Graham 1994). The secretion contains, besides a minute proportion of the precursors for the purple pigment, mucus, proteins, toxins, and narcotizing nar·co·tize  
tr.v. nar·co·tized, nar·co·tiz·ing, nar·co·tiz·es
1. To place under the influence of a narcotic.

2. To put to sleep; lull.

3. To dull; deaden.
 compounds. However, the hypobranchial gland of Plicopurpura pansa (Gould, 1853) is an exception among the muricids. It is so active that the snails can be stimulated periodically to expulse the secretion without being harmed (Rios-Jara et al. 1994; Michel-Morfin & Chavez, 2000).

The carnivorous car·niv·o·rous  
adj.
1. Of or relating to carnivores.

2. Flesh-eating or predatory: a carnivorous bird.

3.
, gonochoristic marine purple snail P. pansa inhabits intertidal in·ter·tid·al  
adj.
Of or being the region between the high tide mark and the low tide mark.



in
 rocky shores exposed to the high impact waves of the open sea. The range of distribution of P. pansa extends from the northwest coast of Mexico (Baja California Sur Baja California Sur (sr), state (1990 pop. 317,764), 27,571 sq mi (71,428 sq km), NW Mexico, on the Baja California peninsula. La Paz is the capital. ) (Clench 1947, Keen 1971) to northern Peru (Pena 1970, Paredes et al. 1999). The snail is not too small (shell length: average about 30 mm, but can reach a total shell length of 90 mm), and at low tides it is relatively easily gathered.

As in antiquity in the Old World, the use of muricids to obtain Tyrian purple for dyeing led also on the Pacific coast of Central America Central America, narrow, southernmost region (c.202,200 sq mi/523,698 sq km) of North America, linked to South America at Colombia. It separates the Caribbean from the Pacific.  and Mexico to a product of high economic value. Here, mainly P. pansa is until now exploited by indigenous communities for Tyrian purple for dyeing cotton yarns and subsequently woven into traditional dresses (Turok 1999). The knowledge about the use of Tyrian purple from P. pansa before Columbus is limited. The first report known to me originates from Fernandez de Oviedo who visited in 1529 the peninsula of Nicoya (Costa Rica Costa Rica (kŏs`tə rē`kə), officially Republic of Costa Rica, republic (2005 est. pop. 4,016,000), 19,575 sq mi (50,700 sq km), Central America. ) and reported the use of "shells or oysters of purple" by natives to dye "mantas, cotton thread, and other things" (Fernandez de Oviedo [not dated], cited by Gerhard 1962). About thirty years later reported Vazquez de Coronado the use of shellfish dyed thread as a minor article of commerce in the province of Quepo in Costa Rica (Vazquez de Coronado 1563).

As Rios-Jara et al. (1994) and Michel-Morfin and Chavez (2000) reported can P. pansa be "milked" periodically to obtain Tyrian purple without harming the animal, however, too frequent milking caused mortalities. It is interesting however, to observe in reports that the exploitation of P. pansa for its pigment caused drastic declines of the snail population (Nuttall 1909, Turok & Acevedo 2000). Yes, in Nicoya, because of the increasing demands for purple dyed material the declining snail population led in 1760 to an uprising of the "indios" against the Spanish Alcalde alcalde (ălkăl`dē, Span. älkäl`dā) [Span., from Arab.,=the judge], Spanish official title, in existence at least from the 11th cent. Since the late 19th cent.  (Fernandez-Guardia 1938, Jinesta 1940). In recent years with the increasing interest in natural colors, the commercial exploitation of P. pansa for dyeing kimonos with "Tyrian Purple" had reached in Mexico such levels as to threaten the survival of the species. In 1988 P. pansa had to be declared by the Mexican government a species under special protection and the government permitted only Indian communities to follow the traditional exploitation of P. pansa for its pigment (Anonymous 1988, 1994).

But why did the populations of P. pansa decline so drastically? Is it because of too frequent "milking" without giving the snails time to recuperate re·cu·per·ate
v.
To return to health or strength; recover.
? Is this causing mortalities or is it the stress from removing the snails from the rocks? Are there other reasons responsible? Low recovery rates of marked purple snails from areas of high wave actions point to the additional possibility that they are just washed away after milking by the waves (Ramirez-Rodriguez & Naegel 2003).

Information about the impact of periodically "milking" of the snails and survival, the percentage of animals showing secretion, and the time for recovery of the hypobranchial gland needed after being "milked" is a prerequisite for the assessment, planning, and management of the potential exploitation of P. pansa for its "Tyrian purple."

MATERIALS AND METHODS

About 300 adult snails were collected, during low tides between October and December 1999, from exposed low intertidal rocks at Playa playa
 or pan or flat or dry lake

Flat-bottomed depression that is periodically covered by water. Playas occur in interior desert basins and adjacent to coasts in arid and semiarid regions.
 Cerritos (23[degrees]19'54"N, 110[degrees]10'38"W) about 120 km southwest of La Paz, Baja California Sur
For other uses, see La Paz (disambiguation).
La Paz is the capital of the Mexican state of Baja California Sur and an important regional commercial center. The city had a 2005 census population of 189,176 persons.
, Mexico. The snails were transferred to the laboratory (CICIMAR, La Paz La Paz, city, Bolivia
La Paz (lä päs), city (1992 pop. 713,378), W Bolivia, administrative capital (since 1898) and largest city of Bolivia. The legal capital is Sucre.
) and kept in inverted inverted

reverse in position, direction or order.


inverted L block
a pattern of local filtration anesthesia commonly used in laparotomy in the ox.
 glass carboys with cut-off cut-off Anesthesiology The point at which elongation of the carbon chain of the 1-alkanol family of anesthetics results in a precipitous drop in the anesthetic potential of these agents–eg, at > 12 carbons in length, there is little anesthetic activity,  bottoms, each filled with 10 L of seawater seawater

Water that makes up the oceans and seas. Seawater is a complex mixture of 96.5% water, 2.5% salts, and small amounts of other substances. Much of the world's magnesium is recovered from seawater, as are large quantities of bromine.
 (salinity: 30 [per thousand] to 34 [per thousand] daily water exchange; temperature range: 21[degrees]C to 23[degrees]C; 12 h light/dark). Sex determination was accomplished by visual inspection for a penis. During the experimental period the animals were fed to satiation sa·ti·a·tion
n.
The state produced by having had a specific need, such as hunger or thirst, fulfilled.



sa
 with daily additions of squid.

For the here presented experiments (August 20 to November 25, 2003) 110 sexed and tagged animals were distributed randomly in 7 glass carboys. For the daily milking experiments one carboy was used with 8 males (length: 29.05 [+ or -] SD 4.31 mm) and 8 females (length: 30.97 [+ or -] SD 3.6 mm), one carboy with 15 makes (length: 26.8 [+ or -] SD 3.15 mm), and another with 15 females (length: 33.04 [+ or -] SD 6.62 mm); for the weekly, biweekly, three-weekly, and monthly milking experiments, the remaining four carboys contained 8 males and 8 females each. The size of all the 55 males ranged from 21.92-36.0 mm (27.9 [+ or -] SD 3.55) and of the females from 23.07-56.01 mm (32.7 [+ or -] SD 6.67). In the carboy with the 15 males for daily milking, on the 22nd day of culture one snail fell to the floor, slightly damaged its shell and died on the 27th day because of the damage. The same accident happened with one female snail, which dropped to the floor and damaged its shell after 20 days in culture and died after 39 days. It was recorded which marked animal expulsed secretion after slight mechanical pressure on the operculum operculum /oper·cu·lum/ (o-per´ku-lum) pl. oper´cula   [L.]
1. a lid or covering.

2. the folds of pallium from the frontal, parietal, and temporal lobes of the cerebrum overlying the insula.
. To determine the effect of daily removing the animals from the carboy without "milking" them, one carboy with 19 animals (8 males: 27.61 [+ or -] SD 2.78 and 9 females: 31.76 [+ or -] S.D. 5.35) was additionally installed.

To compare the proportion of animals that expulsed secretions from the hypobranchial gland under natural conditions, laboratory snails expulsion experiments were undertaken at Playa Cerritos in September 2003 (total number of snails collected 219), in October 2003 (total number of snails collected: 157), and in May 2004 (total number of snails: 455). In September and October 2003 additionally, the sex and total length of the animals were determined.

RESULTS

At the beginning of the experiments attempts were made to milk all 110 animals (55 males and 55 females). It was successful with only 30% (16 males and 17 females). No relation could be found between the size and sex of the animals and the frequency of secretions (Fig. 1).

[FIGURE 1 OMITTED]

From 23 male and 23 female snails, attempts were made to obtain daily secretions for a period of 98 days. During this period 1 male and 1 female died because of handling accidents and additionally 3 males and 2 females for unknown reasons.

At the beginning of the experiment (day 1), of the 46 animals, secretion was expulsed from only 9 (39.1%) males and 8 females (34.8%). The occurrence of secretions declined drastically during the experiment, and during the last weeks only few expulsions were noted (Fig. 2). No correlation could be noted between the sex of the snails and the decline of secretions

[FIGURE 2 OMITTED]

One interesting finding was, that the total number of secretions during the 98 days of the experiment was nearly double (122) from the surviving 20 females than that from the 19 males (68). During the test period each male excreted on average 3.6 times and each female 6 times. However, great individual differences could be observed by counting the total number of expulsions from each snail during the test period. During the 98 days five males did not expulse at all and one 11 times, one female 11 times, four females 6 times, and one not even once. No relation could be found between the size of the animals and the frequency of expulsion (Fig. 3). The smallest male with a shell length of 22.4 mm expulsed 6 times and the second largest one with a length of 32.48 mm 11 times. A similar picture can be seen with the female snails: the second smallest with a shell length of 24.62 mm expulsed 11 times and the second largest with a shell length of 40.16 mm 9 times. From these results it can be concluded that the frequency of expulsion is a characteristic of the individual snail and is not related to sex or size, although female snails expulsed more frequently than males.

[FIGURE 3 OMITTED]

To compare the proportion of expulsions of free-living animals with the laboratory snails, in September and October 2003, and in May 2004 experiments were undertaken at Playa Cerritos. In September 2003, 219 animals were collected; in October 157 animals; and in May 2004, 455 snails. Additionally in September and October 2003 the sex and total length of the collected animals were determined.

From the 831 animals collected 463 expulsed secretion (55.7%). From 376 sexed animals, 223 were males from which 123 expulsed (55.2%) and from the 153 females, 76 (49.7%) expulsed.

We tried to determine the effects of weekly, fortnightly fort·night·ly  
adj.
Happening or appearing once in or every two weeks.

adv.
Once in a fortnight.

n. pl. fort·night·lies
A publication issued once every two weeks.
, three-weekly, and monthly milkings on the frequency of secretion. At the beginning of the weekly experiment three expulsions were observed each from the eight male and eight female snails (31.4%). As in the experiment with the daily "milkings" the frequency of expulsions decreased drastically during the experiment. At the end of the experiment only one male and one female excreted (12.5%) (Fig. 4). The total number of secretions during the test period of 14 wk was significantly higher with the eight females (39) than with the eight males (27), but no relation between the size of the animals and the frequency of expulsion could be found. All animals survived the experimental period.

[FIGURE 4 OMITTED]

A similar situation could be found in the fortnightly "milking" experiment. At the start the experiment only two (25%) excretions were obtained from the eight males, and from the eight females only three (37.5%). The occurrence of expulsions declined during the test period. As an average, only 18% of the animals expulsed (Fig. 5). The number of the total expulsions during the test period was nearly double (13) by the female snails than by the males (7). Also in this experiment great individual differences were found, which were not related to the size, between the animals and the frequency of expulsions. From the eight males three animals didn't expulse during the test period, 4 animals once, and one animal three times. From the eight females four didn't expulse, two animals twice, one animal four times, and one animal even five times.

[FIGURE 5 OMITTED]

"Milking" every 3 wk and monthly had a different effect on the frequency of expulsions as reported before (Fig. 6). At the beginning of the three-weekly experiment with the eight females three expulsed, but with the eight males not one; however, at the end of the experiment the number of expulsions by males (12) and females (13) was nearly the same. Also here large differences could be observed between the individual frequency of expulsions. From the eight males one did not expulse at all during the experiment, and two expulsed three times. From the eight females two did not expulse at all, and one animal expulsed four times. As an average 31.5% of the animals expulsed during the test period and no decline in the frequency of expulsions could be found.

[FIGURE 6 OMITTED]

The effect of the period of recuperation recuperation /re·cu·per·a·tion/ (-koo?per-a´shun) recovery of health and strength.
recuperation,
n the process of recovering health, strength, and mental and emotional vigor.
 between "milking" and the mortality: All animals in the weekly, fortnightly, three-weekly, and monthly experiment survived the "milking" during the experimental period of 98 days. However, a few mortalities were observed in the daily "milking" experiment. During handling of the snails on the 19th day of the experiment one female and one male dropped to the concrete floor and their shells were slightly damaged. The female died after 8 and the male after the l0 days. For unknown reasons another three males died on day 59, 62, and 70; and two females on day 69 and 76. The "milking" was not the reason for the mortalities. For the males the survival rate was 82.6%, for the females 87%.

To determine whether the daily removal of snails is causing mortalities, snails were removed daily from a carboy without "milking " them, and after a few minutes they were returned into the water again. During the test period of 98 days not one snail died because of the daily removal. This result shows that the removal of snails did not lead to mortalities.

DISCUSSION

One of the main functions of the hypobranchial gland is, according to according to
prep.
1. As stated or indicated by; on the authority of: according to historians.

2. In keeping with: according to instructions.

3.
 Fretter and Graham, (1994) to produce secretion for trapping and cementing particulate matter particulate matter
n. Abbr. PM
Material suspended in the air in the form of minute solid particles or liquid droplets, especially when considered as an atmospheric pollutant.

Noun 1.
 sucked into the mantle cavity in the respiratory water current, prior to its expulsion. The gland is a thin band-like, inconspicuous in·con·spic·u·ous  
adj.
Not readily noticeable.



incon·spic
 glandular epithelium glandular epithelium
n.
Epithelium made up of cells that produce secretions.
 consisting of large (150-[micro]m) cells located at the superior part of the internal mantle cavity without excretory ducts, or organs to store the secretion. Like the salivary gland salivary gland

Any of the organs that secrete saliva. Three pairs of major glands secrete saliva into the mouth through distinct ducts: the parotid glands (the largest), between the ear and the back of the lower jaw; the submaxillary glands, along the side of the lower jaw;
 it secretes mucus continuously and can rapidly increase the secretion when stimulated. Together with the mucus minute amounts of the precursors of Tyrian purple are excreted, which lead on the surface of unwashed carboys to a purple coloration col·or·a·tion  
n.
1. Arrangement of colors.

2. The sum of the beliefs or principles of a person, group, or institution.
 of the mucus (unpublished personal observations). When the animal contracts vigorously the cells of the hypobranchial gland burst open with their contents dispersed into the mantle cavity, where it mixes with the mucus and seawater (Lacaze-Duthiers 1859). Because of the few muscle fibers around the hypobranchial gland of muricids, the likelihood of mechanical stimuli is uncertain. The secretion also could be stimulated by neurosecretory neurosecretory

pertaining to or emanating from the secretory activities of nerve cells.


neurosecretory bodies
the form in which neurosecretions are passed along axons to release them into the blood.
 activities because of the presence of neurosensory neu·ro·sen·so·ry
adj.
Of or relating to the sensory activity or functions of the nervous system.
 cells forming the hypobranchial nerve (Srilakshmi 1991), although there is no evidence available about a connection between the nerve and the secretory secretory /se·cre·to·ry/ (se-kre´tah-re) (se´kre-tor?e) pertaining to secretion or affecting the secretions.

se·cre·to·ry
adj.
Relating to or performing secretion.
 cells. In a histologic his·tol·o·gy  
n. pl. his·tol·o·gies
1. The anatomical study of the microscopic structure of animal and plant tissues.

2. The microscopic structure of tissue.
 study, Bolognani-Fantin and Ottaviani (1981) observed that in the hypobranchial gland of M. brandaris there were six different cell types. Roller et al. (1995) studying by means of light and electron microscopy electron microscopy

Technique that allows examination of samples too small to be seen with a light microscope. Electron beams have much smaller wavelengths than visible light and hence higher resolving power.
 the hypobranchial gland of the estuarine es·tu·a·rine  
adj.
1. Of, relating to, or found in an estuary.

2. Geology Formed or deposited in an estuary.

Adj. 1. estuarine - of or relating to or found in estuaries
estuarial
 snail Stramonita (= Thais) haemastoma canaliculata, described eight distinct cell types that are randomly distributed in the gland. Bolognani-Fantin and Otta Ottaviani (1981) questioned whether the great number of different cell types with different functions a characteristic of the hypobranchial gland of "Tyrian Purple," were producing muricids.

The volume of secretion from the hypobranchial gland released into the mantle cavity depends not only on the size but also on environmental and the physiologic conditions of the animals (Born 1936, Fouquet 1970). A large proportion of the volume of secretion obtainable from P. pansa after stimulation originates from the stored liquids of the mantle cavity. The concentration of the pigment precursors in the secretion of P. pansa could not, because of analytical difficulties, be quantitatively determined. Friedlander (1909) isolated 1.4 g of the pure pigment from 12,000 hypobranchial glands of Murex mu·rex  
n. pl. mu·ri·ces or mu·rex·es
Any of various marine gastropods of the genus Murex common in tropical seas and having rough spiny shells, especially M. trunculus, the source of Tyrian purple.
 brandaris and showed that it was 6,6'dibromindigo. Using high performance liquid chromatography High-performance liquid chromatography (HPLC) is a form of column chromatography used frequently in biochemistry and analytical chemistry. It is also sometimes referred to as high-pressure liquid chromatography.  (HPLC HPLC high-performance liquid chromatography.

HPLC

high performance liquid chromatography.

HPLC High-performance liquid chromatography Lab instrumentation A highly sensitive analytic method in which analytes are placed
) for the determination of the chemical composition of the final dye from P. pansa, it could be demonstrated that it consisted out of 90% 6,6'-dibromindigo, 9% monobromoindigo, and 1% dibromoindirubin (Withnall et al. 2003).

Repetitive stimulation to obtain the secretion from the hypobranchial gland without allowing a recuperation time of at least three weeks, results not only in a diminishing expulsed volume (Nuttall 1909, Michel-Morfin & Chavez 2000), but also, as demonstrated in this study, in an increase in the proportion of snails that didn't excrete excrete /ex·crete/ (eks-kret´) to throw off or eliminate by a normal discharge, such as waste matter.

ex·crete
v.
To eliminate waste material from the body.
 at all. Michel-Morfin and Chavez (2000) reported an increase of snail mortalities with decreasing time of recuperation between the milkings. This is not consistent with the results presented here. The attempt to obtain daily secretions over a period of 98 days resulted in minor mortalities, mainly because of mistakes in handling the animals.

Differences between results concerning the proportion of expulsions obtained in the laboratory and the values obtained under natural conditions could be explained by the completely different environmental and physiologic conditions. The mucus production from the hypobranchial gland facilitates the locomotion locomotion

Any of various animal movements that result in progression from one place to another. Locomotion is classified as either appendicular (accomplished by special appendages) or axial (achieved by changing the body shape).
 of the animals. The snails in the field have to produce much more mucus because of the rough surface of the rocks than the animals cultured in the smooth glass carboys. Additionally, the snails in the laboratory were fed only squid, a food completely different compared with those from the intertidal zone, where the snails fed on other snails.

In mark and recapture Mark and recapture is a method commonly used in ecology to estimate population size and population vital rates (i.e., survival, movement, and growth). This method is most valuable when a researcher fails to detect all individuals present within a population of interest every time  experiments at Playa Cerritos the average percentage of first recapture was only 1%, probably because the marked snails deposited among the rocks where they were found did not adhere fast enough, and they were washed away to unsuitable areas by the high wave action. In contrast at a location with low impact waves, the average percentage of first recapture of marked snails was 22% (Ramirez-Rodriguez & Naegel 2003). The main reason for the declining snail P. pansa populations in the areas if "Tyrian Purple" production seems to be the likelihood of being washed away by high impact waves after "milking." For this reason the snails should never be removed from the rocks, and dyeing materials with Tyrian purple should only, in exceptional cases, be permitted to the indigenous people to allow the survival of an tradition.

ACKNOWLEDGMENTS

The author expresses his thanks to C.A. Aguilar-Cruz (CICIMAR) for his technical assistance during the experiments, A. Rosales-M. and I. Castanon-E (U. Nayarit) for their help in determining the sex of the snails, and to J. Lopez-Rocha (CICIMAR), M. Delgado-T. (Tecnologico Culiacan) and O. Armenariz (CIBNOR) for improving the drawings. Special thanks to Chris Cooksey (London) for making suggestions to improve earlier versions of this manuscript and to the two anonymous reviewers for their comments. This work was supported by a grant from PIFI to C. A. Aguilar-Cruz and from COFAA and EDI (Electronic Data Interchange) The electronic communication of business transactions, such as orders, confirmations and invoices, between organizations. Third parties provide EDI services that enable organizations with different equipment to connect.  to the author.

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Presence of hemorrhages in the skin, often associated with bleeding from natural cavities and in tissues. Major causes include damage to small artery walls (as in vitamin deficiency or allergic reaction) and platelet deficiency (in association with such disorders as
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adj.
1. Of special importance or significance; exceptional: an occasion of especial joy.

2.
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n.
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Baja California (Span.: bä`hä kälēfōr`nyä), state (1990 pop. 1,660,855), 27,628 sq mi (71,576 sq km), NW Mexico, on the Baja California peninsula. Mexicali is the capital.
 to Peru. Stanford: Stanford University Stanford University, at Stanford, Calif.; coeducational; chartered 1885, opened 1891 as Leland Stanford Junior Univ. (still the legal name). The original campus was designed by Frederick Law Olmsted. David Starr Jordan was its first president.  Press. 1064 pp.

Lacaze-Duthiers, H. 1859. Memoire sur la pourpre. Annales de Sciences Naturelles. Pattie Zoologique 4e serie. Zoologie. 12:5-84.

Michel-Morfin, J. E. & E. A. Chavez. 2000. Effect of repetitive dye extraction over yield and survival rate of the purple snail Plicopurpura pansa (Gould, 1853). J. Shellfish Res. 19(2):913-917.

Nuttall, Z. 1909. A curious survival in Mexico of the use of the purpura shellfish for dying. In: Putnam Anniversary Volume. Cedar Rapids Cedar Rapids, city (1990 pop. 108,751), seat of Linn co., E central Iowa, on the Cedar River; inc. as a city 1856. The second largest city in Iowa, it is named for the surging rapids in the river. , Iowa: The Torch Press. pp. 366-384.

Paredes, C., P. Huaman, F. Cardoso, R. Vivar & V. Vera. 1999. Estado actual del conocimiento de los moluscos acuaticos en el Peril. Revista Peruana Biologia. 6(1):5-47.

Pena, G. G. M. 1970. Zonas de distribucion de gaster gaster /gas·ter/ (gas´ter) [Gr.] stomach.

gas·ter
n.
The stomach.



gaster

[Gr.] see stomach.
6 podos marinas del Peril. Anales Cientrlicos de la Universidad Agraria 8:153-170.

Ramirez-Rodrfguez, M. & L. C. A. Naegel. 2003. Growth of the purple snail Plicopurpura pansa in Baja California Sur, Mexico. Rev. Ciencias Marinas. 29(3):283-290.

Rios-Jara, E., H. G. Leon-Alvarez, L. Lizarraga-Chavez & J. E. Michel-Morfin. 1994. Produccion y tiempo de recuperacion del tinte Tinte () is a town in the Dutch province of South Holland. It is a part of the municipality of Westvoorne, and lies about 6 km north of Hellevoetsluis.  de Plieopurpura patula pansa (Neogastropoda: Muricidae) en Jalisco, Mexico. Rev. Biol. Trop. 42(3):537-545.

Roller, R. A., J. D. Rickett & W. B. Stickle stick·le  
intr.v. stick·led, stick·ling, stick·les
1. To argue or contend stubbornly, especially about trivial or petty points.

2. To have or raise objections; scruple.
. 1995. The hypobranchial gland of the estuarine snail Stramonita haemastoma canaliculata (Gray) (Prosobranchia: Muricidae): a light and electron microscopical study. American Malacological Bulletin 112:177-190.

Srilakshmi, G. 1991. The hypobranchial gland of Morula morula /mor·u·la/ (mor´u-lah)
1. the solid mass of blastomeres formed by cleavage of a zygote.

2. an inclusion body seen in circulating leukocytes in ehrlichiosis.
 granulata (Gastropoda: Prosobranchia). J. Mar. biol. Ass. UK. 71:623-634.

Turok, M. 1999. Purpura pansa: Una historia de tintes y caracoles, http:// www.conabio.gob.mx/institucion/conabio_espanol/doctos/purpura.html.

Turok, M. & J. Acevedo. 2000. Protection of the colorful Mixteca and Nahoa indigenous dye traditions in Mexico: the saga of the Plicopurpura pansa snail. In: Use of incentive measures for conservation and sustainable use Sustainable use is the use of resources at a rate which will meet the needs of the present without impairing the ability of future generations to meet their needs. The concept was notably put forth by the Brundtland Commission in 1987. See also
  • http://www.iucn.
 of biological diversity. The Hague: United Nations Environmental Program (UNEP UNEP United Nations Environment Program(me)
UNEP Unbundled Network Element Platform
UNEP University of Northeastern Philippines
). pp. 131-157.

Vazquez de Coronado, J. 1563. Juan Vazquez Juan Vazquez may refer to:
  • Juan Bautista Vázquez, a Spanish sculptor and painter of the Renaissance period.
  • Juan Vázquez de Mella, a Spanish politician known for his rhetorical power.
  • Juan T. Vazquez Martin, a Cuban abstract painter who lives and works in Havana.
 de Coronado al muy illustre Senor Don Juan Don Juan (dŏn wän, j`ən, Span. dōn hwän), legendary profligate.  Martinez de Landecho, del Consejo de Su Mag y su Presidente y Governador del distrito de la Audiencia de los confines pp. 227-229 In: D.M.M De Peralta, 1883. Costa-Rica, Nicaragua y Panama en el siglo El Siglo is a Chilean weekly that is the official organ of the Chilean Communist Party's Central Committee. It was founded as newspaper on August 31, 1940.

On July 14, 1948 it was closed down as consequence of the anti-communist Defense of Democracy Law.
 XVI. Su historia y sus lfmites segtin los documentos del Archivo de Indias de Sevilla, del de Simancas etc. recogidos y publicados con notas y aclaraciones historicas y geograficas. Madrid and Paris.

Withnall, R., D. Patel, C. Cooksey & L. Naegel. 2003. Chemical studies of Purpura pansa. Dyes in History and Archaeology. 19:108-117.

LUDWIG C. A. NAEGEL *

Centro Interdisciplinario de Ciencias Marinas, Instituto Politecnico Nacional (CICIMAR/IPN) Apdo. Postal 592 La Paz, B.C.S. 23000 Mexico

* E-mail: lnaegel@ipn.mx
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Author:Naegel, Ludwig C.A.
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Date:Jan 1, 2005
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