Periclimenes+brevicarpalis,+five-spot+anemone+shrimp

The wonderful ocean harbours magnificent organisms, one such example is the //Periclimenes brevicarpalis// you see here.

TL;DR (Too long dont read) option: have a brief overview of this shrimp through a video! media type="custom" key="28809516"

Video adapted from Youtube. Creative commons.

=1. Introduction= This page will introduce you to the //Periclimenes// //brevicarpalis//, otherwise locally known as the five-spot anemone shrimp toc (which also goes by many other names as you will see later). Anemone shrimp refers to a broad list of species inhabiting

[|cnidarian] (anemone) hosts. Its local common name, five-spot anemone shrimp may have been given due to its five orange spots outlined in black colour on its [|caudal] fin (aka tail). This shrimp is known for its transparent body and association with anemones and can be encountered during inter-tidal walks on our local shores in areas such as Chek Jawa or Changi Beach Park.

For the hobbyist, unfamiliar words will be explained in common terms. Otherwise a link will be provided so you will be brought to a page which further explains the technical term. For the scientist, this page will hopefully cater to your scientific desire. Information on this page can be for everyone, however towards the end of the page it might get more technical. Please navigate to a section of your interest with the navigation panel on the right! Enjoy your stay! 

=2. Biology =

2.1 Symbiotic interactions with invertebrate hosts
 Adults of the //P. brevicarpalis// have been found inhabiting sea-anemones belonging to the genus //[|Thalassianthus], [|Stichodactyla], [|Entacmaea], [|Heteractis], [|Parasicyonis]//, and rarely on the coral //[|Acropora].// The anemones may also host other shrimps or fishes such as [|Amphiprion]and also [|porcellanid]crabs. //P. brevicarpalis//, being a generalist, can be found on at least 8 different tropical anemone host. A single anemone often hosts a pair of //P. brevicarpalis//, and often one of each sex.  On the right is a pair of //P. brevicarpalis// inhabiting the Haddon's carpet anemone (//[|Stichodactyla haddoni])//.  According to Holthuis (1952), the youngs of the species are found among weeds on high sea. By this method, the larvae can travel great distances before settlement, helping in dispersal.
 * [[image:taxo4254/magnificent.jpg width="492" height="396"]] || [[image:taxo4254/giantcarpet.jpg width="492" height="396"]] ||
 * //P. brevicarpalis// on a Magnificent anemone (//Heteractis magnifica//) on the left and on a Giant carpet anemone (//Stichodactyla gigantea//) on the right. Other commensals such as the //Amphiprion// fish (aka Clownfish) also shares its home, as shown on the right. Images by [|Ria Tan]. ||

__**2.1.1 A toxic relationship?**__
The relationship between these shrimps and their host is at present known to be [|symbiotic](interdependent), since studies have yet to prove if cnidarian hosts confers protection to the shrimp or if the shrimps are harmful to their host. However, the //P. brevicarpalis// has been documented to clip off the tentacles of the host therefore its relationship may be termed as [|parasitic](causing harm), although no serious damage can be found on the anemone [|in-situ] (in the field). On the contrary, similar host associated shrimps of other genus have been found to benefit (in the case of //[|Alpheus armatus]//) the host by protecting it from predation from [|annelid] worms instead. This may be an instance of a love-hate relationship (except being immobile, the anemone cannot run away from a toxic relationship).

__**2.1.2 Just a home or more?**__
Cnidarian hosts are often perceived as protection from predators for the shrimp in the wild. However, in laboratory experiments, //P. brevicarpalis// had reduced growth, longevity and fecundity without the presence of a host (despite being fed with other food sources). //P. brevicarpalis// larvae were also unable to develop ovaries in the absence of a host for settlement. This may suggest a more obligatory relationship of the host association in the //P. brevicarpalis// than only for protection from predation (aka being eaten up).

2.2 Feeding habits
<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14.3px;">//Periclimenes brevicarpalis// being [|commensals] (a relationship where one feeds off the other, much like in some romantic human relationships), feeds on the leftover food particles of the host. Although when food is scarce, this shrimp also feeds on the host it associates with. In the day, they can be found most often on the periphery of [|oral disc] (top part of the anemone you most often see) on the anemone, clipping and eating its outer tentacles. In the wild, this behaviour has not been observed to be especially harmful to the anemone as they often still retain their long and numerous tentacles.

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14.3px;"> //Periclimenes brevicarpalis//, although graceful and magnificent looking, also feed on each other. This behaviour is most noticeable after molting and in ex-situ (out of field) conditions when they were reared together in a tank. Cannibalism may not occur in the wild due to the fact that these shrimps are often only found in pairs on a single anemone. Or they may have already eaten the previous residents, who knows!

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 16.9px; text-align: justify;">2.3 Reproduction
<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14.3px;">[|Ovigerous]females can carry up to 4000 oval shaped eggs until they hatch. However insufficient space for accomodation of the clutch, as well as environmental stressors may induce brood loss.The association with an anemone may reduce brood loss during embryogenesis as it provides favourable conditions for embryo development. This species has been observed to release approximately 300 youngs at once. The larvae of this species has a long pelagic larvae duration (PLD) and may settle only 35 days after hatching. <span style="font-family: Arial,Helvetica,sans-serif; font-size: 14.3px;"> While other species of Palaemonidae shrimps undergoes mark-time moults (where zoeal stage larvae moults multiple time without change in morphology but only in size) and sometimes skip stages, the development of //P. brevicarpalis// is noted to be much more regular.

2.3.1 Life stages
<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14.3px;">This table summarises the different life stages of the //P. brevicarpalis// days after hatching, with illustration for the last zoea stage to the first juvenile stage to show its transformation into an immature adult from a pelagic larvae. <span style="font-family: Arial,Helvetica,sans-serif; font-size: 14.3px;">Illustrations by Nagai and Shokita, 2003. Pending permission.
 * ~ Stages ||~ 1st [|zoea] ||~ 2nd zoea ||~ 3rd zoea ||~ 4th zoea ||~ 5th zoea ||~ 6th zoea ||~ 7th zoea ||~ 8th zoea ||~ 9th zoea ||~ [|Decapodid]stage ||~ First [|juvenile]stage ||
 * = **Days after hatching** ||= 1-2 ||= 2-4 ||= 4-6 ||= 6-8 ||= 8-10 ||= 10-14 ||= 12-17 ||= 14-20 ||= 16-27 ||= 18-31 ||= 20-35 ||
 * =  ||=   ||=   ||=   ||=   ||=   ||=   ||=   ||=   ||= [[image:9thzoea.JPG width="202" height="151"]] ||= [[image:decapodid.JPG width="215" height="151"]] ||= [[image:firstjuv.JPG width="162" height="107"]] ||

=<span style="font-family: Arial,Helvetica,sans-serif; font-size: 16.9px; text-align: justify;">3. What may threaten the survival of this species? = <span style="font-family: Arial,Helvetica,sans-serif; font-size: 14.3px;">Marine tropical decapod crustaceans are among the most popular invertebrate species in the aquarium trade industry. Due to its attractive colouration and its interesting host association, this species is an ornamental species. Some aquariums also promote them as having a peaceful temperament and being easy to care for, which may attract more buyers, increasing demand and eventually increasing harvesting from the wild. A google search will these shrimps at attractive prices below SGD20 and sometimes even with free shipping. While it is not known whether our local populations are being poached, other countries heavily involved in the aquaculture trade industry may be.

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14.3px;"> Given the lack of understanding for culture techniques of these ornamental species and their obligatory host association, they are often harvested from the wild. This practice not only endangers the organism, but is also likely to impact the ecosystem due to their associative behaviours with other organisms. Sustainable management of the collection of ornamental species such as this will need to be implemented for their conservation. Another sustainable solution is to convince marine aquarium hobbyists to rear shrimps which can be artificially cultured instead.

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=<span style="font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">4. How do you tell if you are looking at the right species? =

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">4.1 How to tell it apart from closely related species
<span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">This species closely resembles //[|Periclimenes inornatus]// and //[|Periclimenes ornatus]//, however it may be distinguished by its size and position of the dorsal and terminal spines of the telson. For example in the //P. brevicarpalis//, dorsolateral spine of the telson is on the proximal half of the telson length while that of //P. ornatus// is on a third of the telson length. <span style="display: block; font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">
 * <span style="display: block; font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">[[image:inornatus.ashx width="377" height="383"]] || <span style="display: block; font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">[[image:brevmle.jpg]] ||
 * //P. inornatus// versus //P. brevicarpalis//, bearing some resemblance to each other. Image by [|Ole Johan Brett] (on the left) and [|Ron Yeo] (right). Permission granted. ||

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">4.2 Detailed description of the organism's appearance
Image by [|Florent Charpin], pending permission. || Image of a male specimen in Dumaguete, Philippines. Image by [|Florent Charpin], Pending permission || <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> While the life colouration and overall size of the two sexes may differ, there are some taxonomic characteristics which maintains in both sexes. Note that this section is catered for those with expertise in shrimp taxonomy, as some of the jargon may require prior knowledge to understand. If you are inexperienced in this area, or are just looking for a general identification method, please move on to section 3.3 on the species' life coloration. If you wish to continue, below is an illustration of basic shrimp anatomy to help you along!
 * <span style="display: block; font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">[[image:periclimenesbrevicarpalis7.jpg]] || <span style="display: block; font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">[[image:periclimenesbrevicarpalis2.jpg]] ||
 * Images of a female specimen in Bali, Indonesia.

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 130%;">

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> For the shrimp taxonomists, here are some of the key characteristics for the identification of the //Periclimenes brevicarpalis// according to a taxonomic key by Chace and Bruce (1993) :

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> For an accessible description of this species, please refer to the original paper by Chace and Bruce.
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">Carapace without supraorbital or postorbital tooth
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">Second pereopod without acute distal tooth on flexor margin merus
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">Posteriormost tooth of dorsal rostral series not widely separated from rest of series
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">Hepatic spine not extending beyond anterior margin of carapace
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">3rd pereopod without denticulate lobe on flexor margin of dactyl
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">Third pereopod with dactyl simple, not biunguiculate
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">Rostrum with midrib nearly horizontal, directed more anteriad than anteroventrad
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">Rostrum with dorsal margin distinctly convex
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">First pereopod with fingers pectinate on opposable margins
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">2nd pereopod with fingers nearly as long as palm, carpus 1.5 times as long as distal width

4.3 Color in life
<span style="display: block; font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;"> <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> Specimens are almost transparent, save for some pigmented areas which differs between the male and female. Original description by Kemp (1922). <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> Ovigerous (bearing eggs) females in life are almost transparent except for pigmented areas as follows:

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">According to Kemp and others, considerable variations in the colour among the individuals are generally found. For example in the above image, the female from Bali, Indonesia does not have a posterodorsal white patch.
 * 1) <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">Each articulated part of all the [|pereopod] (walking limbs) is dark blue.
 * 2) <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">The [|carapace] (head shield) has the pure white markings on either side and on the dorsal (upperside) portion where the gonad is placed.
 * 3) <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">In addition, the eyestalk and the part just behind the [|rostrum] (forwards projecting spikey horn) on the dorsal side of the carapace are also white.
 * 4) <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">The dorsal and lateral sides of the [|abdomen]is covered with two pairs of the large, oval and white patches outlined by black bands laterally and the broad white patch situated on the posterodorsal (upperside and towards the end) side. Lateral patches are not seen in males.
 * 5) <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">On the proximal half of the telson and each of the uropods there is a white, circular and broad ring followed by an orange eye-spot which is bordered by a dark, brown band. This is what gives the species its common name of the 5 spot anemone shrimp!

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> The female of this species grows to about 5cm long, while the males are smaller at 3cm.

<span style="display: block; font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">

=<span style="font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">5. Where can you find them? = <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">This species can be found in the tropical [|Pacific] and [|Indian Oceans], including Australia. This species has been recorded throughout the [|Indo-West] Pacific region (Miyake and Fujino, 1968). <span style="display: block; font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">5.1 Where are they found in Singapore?
<span style="display: block; font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">media type="custom" key="28800962" <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">//Periclimenes brevicarpalis// has been found on local large sea anemones like the Giant carpet anemones (//[|Stichodactyla gigantea]//), Haddon's carpet anemones (//[|Stichodactyla haddoni]//), Magnificent anemones (//[|Heteractis magnifica]//), Leathery anemone (//[|Heteractis crispa]//) and Pizza anemone (//[|Cryptodendrum adhaesivum]//). <span style="display: block; font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">media type="youtube" key="RgHRhwzjJrE" width="560" height="315" <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> A //Periclimenes brevicarpalis// wandering around its apartment located at Cyrene Reefs, right in our backyard!

=<span style="font-family: Arial,Helvetica,sans-serif; font-size: 16.9px; text-align: justify;">6. What do scientists and layman call it? = <span style="font-family: Arial,Helvetica,sans-serif; font-size: 14.3px;">[|Binomial] (a two part Latin name of an organism consisting of genus followed by its specific epithet): //Periclimenes brevicarpalis// (Schenkel) 1902

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14.3px;"> [|Synonyms] (scientific names that applies to a taxon that (now) goes by a different scientific name): //Ancylocaris brevicarpalis// Schenkel, //Periclimenes (Harpilius) brevicarpalis//

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14.3px;"> [|Vernacular] (aka common names) : Five-spot anemone shrimp, Peacock tail anemone shrimp, Glass Anemone Shrimp, pacific clown anemone shrimp, White Patched Anemone Shrimp

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 14.3px;"> Original description (for the taxonomists out there): not available online

=<span style="font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">7. Where is the original specimen of this species? =  Species description and name of this species are given based on this type specimen. These specimens are usually stored in a museum so that scientists all over the world can come look at it, to compare with the specimens they have on hand. <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">The morphological vouchers of //Periclimenes brevicarpalis// are housed at [|Ocean Genome Legacy]

=<span style="font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">8. Phylogenetic information = <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> Warning for the layman: section ahead may get too deep if you have no prior understanding, however may still be an interesting read if you want to expand your knowledge as common layman words are used to the best of my ability, otherwise links are provided for a deeper understanding. Enjoy! <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">//Periclimenes// shrimps may be found in both deep and shallow waters, with taxonomic differences such as number of dorsal telson spine differing in species belonging to each depth. This genus of shrimp has also adapted to associating with many different kinds of host, including Echinodermata (starfish, brittle star, sea urchin, feather star, sea cucumber), Mollusca (gastropods, bivalves, cephalopods etc.), Porifera (sponges), Ascidiacea (e.g. sea squirts), Polychaeta (annelid worms) on top of cnidarians. Symbioses with cnidarians however, was most likely the ancestral state of the host associations in symbiotic shrimps, which explains why association with cnidarians are most numerous.
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Kingdom || <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Animalia ||
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Phylum || <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Arthropoda ||
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Subphylum || <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Crustacea ||
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Superclass || <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Multicrustacea ||
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Class || <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Malacostraca ||
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Subclass || <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Eumalacostraca ||
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Superorder || <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Eucarida ||
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Order || <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Decapoda ||
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Suborder || <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Pleocyemata ||
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Infraorder || <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Caridea ||
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Superfamily || <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Palaemonoidea ||
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Family || <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Palaemonidae ||
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Genus || //<span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Periclimenes // ||
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Species || //<span style="font-family: Arial,Helvetica,sans-serif; font-size: 120%;">Periclimenes brevicarpalis // ||

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> Each species of //Periclimenes// has adapted to a group of specific hosts, developing characteristics such as laterally compressed bodies to fit in narrow spaces within branching corals or having unarmed rostrum (a protrusion at the top of the carapace, usually with dorsal and ventral teeth) to preventing irritating the soft body of the host in the case of mollusk. <span style="display: block; font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;"> <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> Due to the variety of host specific relationships, //Periclimenes// is a species-rich group today. Previous classifications based solely on morphology may be inaccurate due to the diversity of this group as well as [|homoplasy], leading to //Periclimenes// being a [|polyphyletic]group which can be observed in the phylogenetic tree below. <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> The species //Periclimenes brevicarpalis// (Schenkel, 1902) is closely related to //Ancylomenes//. The abdomen of the //P. brevicarpalis// is slightly humpbacked (and this can be observed in the above images of the 9th zoeal stage) which is a characteristic of //Ancylomenes//. For the taxonomists, a few other similar characters between these two genus according to Kou et al (2015) <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> are as follows: <span style="display: block; font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;"> <span style="display: block; font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">subcylindrical body
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">well-developed palaemonoid rostrum
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">non-ovate inferior orbital angle
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">feeble biungulate dactyls of the last three pereopods
 * <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">generally transparent body with mottled spots

=<span style="font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">9. Molecular information = <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;">Mitochondrial cytochrome oxidase I Sequence of voucher specimen: []

=<span style="font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">10.External links = <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> Local information on of //Periclimenes brevicarpalis// on [|Wild Singapore] by Ria Tan <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> Taxonomic details of //Periclimenes brevicarpalis// on [|World Register of Marine Species] (WoRMS) <span style="font-family: Arial,Helvetica,sans-serif; font-size: 110%;"> Compilation of //Periclimenes brevicarpalis// information on [|Encyclopedia of Life] (EOL)

=<span style="font-family: Arial,Helvetica,sans-serif; font-size: 130%; text-align: justify;">11. Literature and References =