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Close up of , a large ammonite (, , , Hoplitaceae, Placenticeratidae) with a shell diameter of upto 50 cm. During the , ammonoids were extremely plentiful and diverse. In the middle background is the of a shell. The carniverous neogastropods radiated during the Cretaceous, triggering an among and other organisms on which they preyed.

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The mollusks are a large and diverse group of soft-bodied unsegmented animals. Nearly 130,000 are known, and some 35,000 species (these latter are without doubt only a small fraction of those that have ever lived). They include many familiar animals, like , , , etc, as well as others not so well known. They range in size from microscopic forms to the giant squid (), and have a long and venerable history appearing during earliest time, if not before.

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All mollusks possess some or all of the following characteristics:

  • a muscular fleshy foot which with gastropods (snails) is used to crawl along with but in cephalopods is modified into tentacles
  • a visceral mass containing the digestive, excretory, and reproductive organs
  • a , usually two folds that enclose the gills or lungs, and also secretes the protective shell
  • a , a tongue-like feeding organ, essentially a sort of built-in saw equipped with rows of microscopic teeth that is incredibly effective at scraping food off hard surfaces or grinding through any substance (some carnivorous snails are able to bore a whole through the shells of other mollusks)
  • a special respiratory gill known as the ctenidium, (plural ctenidia)
  • a shell made of calcium carbonate. This fossilizes easily, which explains the rich fossils record the mollusks possess.

Although not all mollusks share all these characteristics, generally all mollusks have at least some of them.

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Despite sharing these common features, the mollusks are an incredibly diverse group. The following diagram shows the major classes, which as can be seen are all based on variation in this same body plan.

Figure from D. R. Prothero, 1998, Bringing Fossils to Life, McGraw-Hill, p. 277; from Clarkson, 1993 - this image from LAB 3 Chapter 8: Phylum Mollusca and Biological Diversity and Evolution through Time

This diagram presents a stylized relationship between the different molluscan groups, with all evolving from a "hypothetical ancestral mollusk". It is now known that molluscan phylogeny is a lot more complex than this.

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As one of the most successful forms of animal life, the mollusks have conquered almost every habitat and exist in all the oceans (from shallow tidal pools to the deepest trenches), in fresh water, and on land. The only environment they cannot cope with are very dry regions, as their moist skin is easily desiccated. Even so, the greatest diversity of mollusks species is to be found in marine environments.

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When was formulating his system of binomial nomenclature the mollusks were grouped (along with most invertebrates) under , worms. The name Mollusca (from the Latin mollis "soft"), was first used by the great French zoologist Cuvier in 1798 to refer to cephalopods like squids and cuttlefish. It was later extended to include other organisms of this group, such as snails and bivalves.

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The mollusks first appeared during the earliest (), at the very start of radiation of animals. As the Cambrian period progressed, many types appeared that soon died out. Although previously included among the conventional classes of mollusks it is increasingly argued that these represent distinct classes, early experiments in molluscan evolution, or transition forms. However, the exact number of extinct classes remains debatable.

By the period the three main classes - Gastropoda, Bivalvia, and Cephalopoda - had increased greatly in number and diversity. This was a period of major evolutionary diversity for the phylum, and they became common in most marine ecosystems.

The saw bivalves invade , and the first land snails appeared during the . During the Devonian also, the took over from the as the dominant cephalopod group, and these creatures continued to flourish throughout the entire , living alongside species of bivalves and gastropods not very different to modern forms. (Oysters for example were common during the Jurassic and Cretaceous, while scallops date back to the ). By mid Cretaceous most mollusks, like most invertebrates and fish, were essentially like modern forms, except for the cephalopods which were still represented largely by the Mesozoic and . The ammonites were decimated by the K-T extinction event, as were the reef-building (although these were on the decline for some time), but the (soft-bodied squids, octopods etc) continued to evolve quite happily.

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Traditionally seven are recognized: (primitive worm-like shell-less forms), (chitons, at one time included with the Aplacophora in the class ), (primitive ancestral forms), (snails and slugs), (clams, mussels, oysters, etc), (tooth and tusk shells), and (squids, octopus, nautilus, cuttlefish), of which only Gastropoda, Bivalvia, and Cephalopoda have large numbers of species and a strong . However it is becoming increasingly recognized that there were more groups that have since died out.

With the , recent work in molluscan systematics is done via . The following is a recent cladogram showing molluscan relationships.

image and text by Ellen Thomas - from

My own position, oft-stated, is that the Linnaean and cladistic methodologies are complementary rather than contradictory. Cladistics provides a means of drawing the family , while the Linnean hierarchy enables us to construct a useful, if arbitrary, arrangement for cataloging and comparative purposes. The following listing therefore represents a possible tally of molluscan classes (including extinct forms) that are known or have been proposed (note - not all experts agree as to the status of the various Paleozoic minor classes).

  • (= Neomeniomorpha; Aplacophora in part) - no fossil record - very small creeping worm-like soft bodied shell-less forms.
  • (= Chaetodermomorpha; Aplacophora in part) - ? to - very small deep sea worm-like soft bodied burrowing shell-less forms. The Silurian , which resembles both Caudofoveata and Polyplacophora, may be an early form of this lineage.
  • (= Amphineura in part) - Late Cambrian to Recent - The chitons, which have eight plates or "valves" which are overlapping in post-Mesozoic forms. The foot is broad; the head reduced. The soft parts show some segmentation. The animal generally lives attached to rocks in the inter-tidal zone. Appear to be distantly related to the Solenogastres and Caudofoveata (as ancestors/cousins), and the Tryblidiida.
  • (= Monoplacophora sensu stricto, in part) - Late Cambrian to Recent - Creeping cap-shelled forms with paired muscle attachments. The most primitive group. A few dozen deep water recent species remain as "living fossils". As with the Polyplacophora the soft parts show some segmentation. Probably evolved from a soft-bodied ancestor, possibly Polyplacophora relationships.
  • (= Monoplacophora in part) - Early to Middle Cambrian - Small group of little known bivalved forms. There relationship with other molluscan groups is unclear, and it is not agreed whether they qualify as class status. No descendents, ancestors unknown.
  • (= Monoplacophora in part; paraphyletic) - Earliest to Late Cambrian - an ancestral lineage or . Consist of creeping or weakly infaunal forms with cap-shaped or spiral shell. Probably evolved from a soft-bodied ancestor, as the direction of shell coiling is opposite that of the Tryblidiida, which they predate in any case. Generally considered directly or indirectly the ancestors of all conchiferan mollusks other than the Tryblidiida.
  • (= Pelecypoda) - Early Cambrian to Recent - mostly sedentary or burrowing forms with a shell of two valves hinged dorsally. The foot is generally hatchet-shaped (hence the name "pelecypod"); the head lacking; a very diverse group, includes or forms, mostly marine but some freshwater species. Clams oysters, mussels, etc. Generally considered to have evolved from rostroconches, although this would relate them to helcionelloids, if not actually scaphopods and cephalopods, and the weight of current molecular and cladistic evidence indicates the bivalves diverged from the main Conchiferan lineage quite early.
  • - Early Cambrian to Devonian - Small group of superficially snail-like forms, and perhaps snail-like habits, distinguished by lack of torsion. Evolved from helcionelloids, no descendents. It is not even certain if this is a natural or a polyphyletic group.
  • "" in part (= Monoplacophora in part; polyphyletic) - Late Cambrian to Devonian - A probably artificial assemblage of creeping cap- or partly spiral shelled forms with single or paired muscle attachments. The Tryblidiida are usually included here but they are certainly a more basal group. Non-tryblidiidan Tergomya may have evolved probably several times from helcionelloids, although some may be related to the Tryblidiida. Some of the helcionelloid-descended forms would seem to be related to the gastropods.
  • - Late Cambrian to Recent - Diverse group of shelled or shell-less forms, evolved from a spiral-shelled helcionelloid or helcionelloid-descended "tergomyan" ancestor that underwent . The body is usually asymmetrical, with a distinct head, pair of eyes, and one or two pairs of tentacles. The shell is usually spirally-coiled, but may (in the case of limpets) be cap shaped, and in some species is or completely absent. The foot is broad and used to creep along the substrate. Includes marine, freshwater, and terrestrial species (the only mollusks to invade the land). Snails, slugs and their relatives.
  • - Early Cambrian to Late - A Paleozoic group of infaunal forms. The shell consists of two valves joined in adults by an enclosed hinge. Evolved from helcionelloids. The posterior of the shell is usually produced into an elongate tube, presumably for inhalant and exhalent siphons. Originally considered to have given rise to both scaphopods and bivalves, as they share many similarities with both those groups. But relationships among early molluscan classes remain controversial.
  • - Devonian to Recent - small semi-infaunal forms, the tubular shell open at both ends and resembles a miniature elephant's tusk, with one end larger than the other. The foot is conical; there are no gills. Probably evolved from rostroconches. Shown to be related to the cephalopods.
  • (= Tentaculitoidea) - Middle Cambrian to Devonian - very small pelagic forms with tubular shells. They are thought to be related to the cephalopods, and seem to have born a number of arms.
  • - Latest Cambrian to Recent - diverse group of large intelligent predaceous forms, with a large external shell (most Paleozoic and Mesozoic forms, and current ), a vestigial internal shell, or (in the case of the octopus) the shell is lost altogether. The head is large with well-developed eyes, and armed with horny chitinous jaws and many arms or tentacles for grasping prey; the head fused to the foot (hence the name). Evolved from helcionelloids.
  • - Earliest Cambrian to Permian - Enigmatic animals with small conical shells, shell-microstructure indicates close to mollusks but experts do not agree whether they actually were mollusks.

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Mollusca (see below for synonymy)
   |  |  `--
   |  |--
   |  `--+--
   |     `--+--Neopilina galatheae
   |        `--Laevipilina antarctica Warén & Hain 1992
      |  |    |--P. lapworthi
      |  |    `--P. wellsi
      |  `--+--
      |     `--+--Jinonicella Pokorny 1978 [Jinonicellidae]
      |        `--Janospira Fortey & Whittaker 1976
      |             `--J. nodus Fortey & Whittaker 1976
      `--+--Acaenoplax Sutton, Briggs et al. 2001
         |    `--*A. hayae Sutton, Briggs et al. 2001
               `--Caudofoveata [Chaetodermomorpha]
                    |  i. s.: Scutopus ventrolineatus
                         |--Chaetoderma nitidulum
                         |--Lepoderma chiastos (see below for synonymy)
                         `--Falcidens Salvini-Plawen 1969
                              |--F. lipuros
                              `--F. poias Scheltema 1995

Mollusca incertae sedis:

   |--Z. mitriformis Adams 1860
   |--Z. subvitrea (Smith 1879)
   |--Z. validicosta (Habe 1960)
   `--Z. zonata (Gould 1860)
 Margaretes umbilicus
 Kelletia kelletii
 Cumingia tellenoides
 Succinodon (n. d.)
 Harmandia Rochebrune 1882
 Deltoideum delta
 Bilobissa dilatata
 Umbella d’Orbigny 1841
 Zenatia acinaces
 Marcia plana
 Lapparia hebes
 Galeodea senex
 Bullinella enysi
 Gilbertia tertiaria
 Mesalia striolata
 ‘Dimorpha’ Gray 1840 non Jurine 1807
   |--Hypseloconus Berkey 1898
   `--Ozarkoconus Heller 1955
 Throopella [Throopellidae]
 Janella Grateloup 1838
 Procarinaria Perner 1911 [Procarinariidae]
 Placopecten magellanicus
 Hyalocylis striata
 Corolla calceola
 Firoloida desmarestia
 Cuma carinifera
 Lagena wiegmanni
 Ruma sebae
 Lampania zonalis
   |--V. angularis
   `--V. unicolor
 Gladius fusus
 Macha strigillata
 Hiatula diphos
 Trigonella luzonica
 Cryptogramma flexuosa
   |--C. gibba
   `--C. undatina
 Anomalocardia trapezina
 Mycetopus [Mycetopidae]
   `--M. soleniformis

Lepoderma chiastos [incl. Falcidens (Lepoderma) targotegulatus Salvini-Plawen 1992]

Mollusca [Aculifera, Amphineura, Ancrypoda, Archinacellida, Archinacellina, Archinacelloidea, Cyrtosoma, Dermobranchea, Diasoma, Ganglioneura, Glossophora, Helcionellacea, Helcionelloida, Hyperstrophina, Isopleura, Loboconcha, Monoconcha, Monoplacophora, Paragastropoda, Pteropoda, Scenelloidea, Securiconidae, Solenoconchia, Spiculata, Univalvia]

Vallatotheca elegantula (Chapman 1916) [=Helcionopsis elegantulum; incl. Tryblidium (Metoptoma) nycteis Cresswell 1893 non Metoptoma nycteis Billings 1862, Capulus nycteis, Helcionopsis nycteis]

* Type species of generic name indicated

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Domning, D. P. 2001. The earliest known fully quadrupedal sirenian. Nature 413: 625-627.

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Fraser, L. 1866. Communication of a list of mollusks collected by R. Swinhoe, Esq., in Formosa. Proceedings of the Zoological Society of London 1866: 146.

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By Amanda Gardner Psychogenic impotence is relatively uncommon. It is characterized objectively by the presence of good nocturnal and morning erections and negative findings from all other tests. During the interview, a history of highly variable erections that can be totally absent one day but virtually normal the next suggests a psychogenic cause. Virtually 100% of men with severe depression have erectile dysfunction (ED). Sildenafil (Viagra) works well for psychogenic ED; other treatment modalities are also effective because the tissues, nerves, hormone levels, and vasculature are normal. The authors usually recommend a full psychological evaluation in these patients so that the underlying etiology can be identified and treated appropriately rather than just treating the symptom of ED. Therefore, the authors defer treatment of the patient's ED until he has begun psychological testing and therapy for the underlying problem. Agents that are injected directly into the penis exert their relaxant effect directly on the corpora cavernosal smooth muscle. They can be used as single agents or in combination. The most commonly used agents include PGE1, papaverine, and phentolamine. The dose and most effective combination of these agents must be determined for each patient. These medications can be obtained commercially as Caverject (PGE1) or can be formulated according to the physician's request by compounding pharmacies. Patients can be supplied with vials of a single agent or a combination of agents mixed in a single vial. Patients must be instructed in the proper technique for administration. A single intraurethral agent, PGE1, which has been formulated into a small suppository, is commercially available as MUSE. This agent was available prior to the introduction of sildenafil and is still used by a select group of men. Success rate of 20-25% for the Clinical Efficacy Assessment Subcommittee of the American College of Physicians* Restriction of blood flow caused by a vascular disease can occur because of a buildup of cholesterol and other substances that may block blood vessels. It also is known that blood vessels open more widely (relax) or close down to become narrower (constrict), possibly due to signals from cells called endothelial cells that are located inside the lining of these blood vessels. These signals can result in the restriction of blood flow to certain organs even without the vessels' necessarily being clogged by any material. When endothelial function is impaired, constriction that is similar to plugged blood vessels may result. This could be one explanation why ED is so often associated with vascular disease. Results: Almost one in 10 of the respondents (9.1%) reported erectile dysfunction that lasted for at least one month over the previous year. More than a quarter (27.2%) of respondents were current smokers, with 20.9% smoking ? 20 cigarettes per day, and 6.3% smoking > 20 cigarettes per day. Compared with non-smokers, the adjusted odds ratios for erectile dysfunction were 1.24 (95% confidence interval (CI) 1.01 to 1.52, p ?=? 0.04) for those smoking ? 20 cigarettes per day and 1.39 (95% CI 1.05 to 1.83, p ?=? 0.02) for those smoking > 20 cigarettes per day, after adjusting for other confounding factors. Older age, low level of education, and taking medications for cardiovascular disease were also independently and positively associated with erectile dysfunction. In contrast, moderate alcohol consumption (14 drinks per day) significantly reduced the likelihood of having erectile dysfunction. HealthDay Reporter TUESDAY, May 22 (HealthDay News) -- Worried about jet lag? Researchers think they might have just the ticket to perk you up: Viagra. In addition, always follow the directions on your prescription label carefully and ask your doctor or pharmacist to explain any part that you do not understand. Take these drugs exactly as directed. Do not take more or less or take it more often than prescribed by your doctor. Do not take these drugs more than once a day. Many men would rather stick a needle in their eye than use injection therapy for impotence. What you need to know about Cialis, Levitra, and Viagra. Taking Viagra Won't Spur Risky Sexual Behavior Requires refrigeration But the drug didn't help hamsters who underwent a simulation of westward jet travel. Kushida suggested that there may be a role for restless legs drugs in erectile dysfunction treatment, and said the question warrants further research. Pulmonary hypertension -- the incurable condition of continuous high blood pressure in the pulmonary artery located in the right ventricle of the heart -- is one of many serious complications that can strike COPD patients. PAH causes the artery, which is responsible for delivering blood from the heart to the lungs, to work harder than normal. A weakening of the heart muscle can ensue over time, increasing the risk of heart failure and even death. Out of 117 men who were suitable for taking sildenafil, the acceptance of sildenafil as a treatment option is high, 98.3% (115/117). Sildenafil was successful in treating ED in 91.3% (84/92) of this study population. At the end of the study period, 79.7% (59/74) of men wanted to continue treatment with sildenafil. The side effect profile of sildenafil in this study would appear to be different from previous published data [6] in that the incidence of headache, flushing, dyspepsia and abnormal vision is higher, but in the absence of a control group the significance of this is uncertain. It must be noted that 13% of men who reported side effects at Visit 2 reported complete lack of them at Visit 3. It is of interest that there is a 2% incidence each of thirst and 'dazed feeling' (reported by patients as a 'muzzy head'). The physiology underlying this is unclear, although it could be related to PDE inhibition in the brain, where the various PDE isoenzymes are in abundance [11]. Drugstore Here are some questions to ask your doctor (e.g., urologist) about impotence. Print this page, check off the questions you would like answered, and bring it with you to your doctor's appointment. The more you and your partner know about erectile dysfunction, the easier it will be to develop and follow a successful treatment plan.

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