رخويات

رخويات
Temporal range: الكامبري المتأخر–الحاضر، 535–0 Ma[1]
تنوع الرخويات. باتجاه عقارب الساعة من أعلى اليسار: Ruditapes philippinarum, Littorina littorea, Dentalium sp., Tonicella lineata, Nautilus pompilius, and Amphioctopus fangsiao
التصنيف العلمي e
أصنوفة غير معروفة (أصلحها): الرخويات
تحت شعبة، أجناس غير مصنفة، وصفوف
Diversity[4]
85.000 نوع حي متعرف عليه.
Cornu aspersum (formerly Helix aspersa)—a common land snail
Shell of Marisa cornuarietis, a freshwater snail

الرخويات (Mollusca، هي شعبة من الحيوانات اللافقارية أولية الفم، والتي يُعرف أعضاؤها بالرخويات[ب] (/ˈmɒləsks/). تم التعرف على 86.600 نوع موجود من الرخويات[6]، مما يجعلها ثاني أكبر شعبة حيوانية بعد مفصليات الأرجل. ويقدر عدد الأنواع الأحفورية الإضافية ما بين 60.000 و100.000،[7] ونسبة الأنواع الموجودة غير الموصوفة عالية جداً. لا تزال العديد من الأصنوفات غير مدروسة بشكل جيد.[8]

الرخويات هي أكبر شعبة بحرية، وتضم حوالي 23% من جميع العضيات البحرية المسماة. وهي متنوعة للغاية، ليس فقط في الحجم والبنية التشريحية، لكن أيضاً في السلوك والموئل، حيث أن العديد من المجموعات هي رخويات مياه عذبة وحتى أنواع برية. تنقسم الشعبة عادةً إلى 7 أو 8 صفوف تصنيفية،[9] منها اثنان بالكامل منقرضان. الرأسقدميات، مثل السبيط، الحبار، والأخطبوط، هي من بين أكثر اللافقاريات تطوراً من الناحية العصبية — وإما الحبار العملاق أو الحبار الضخم هو أكبر أنواع اللافقاريات الموجودة المعروفة. تعد البطن قدميات (الحلزونات، والبزاقات| والحلزونات البحرية) الفئة الأكثر تنوعاً وتمثل 80% من إجمالي أنواع الرخويات المصنفة.

السمات الأربعة الأكثر عالمية التي تحدد الرخويات الحديثة هي جسمها الأملس الذي يتكون بالكامل تقريباً من عضلة، وشاح مع تجويف كبير يستخدم للتنفس والإخراج، ووجود مبرد (باستثناء ذوات المصراعين)، وبنية الجهاز العصبي. بخلاف هذه العناصر المشتركة، تعبر الرخويات عن تنوع شكلي كبير، لذلك تعتمد العديد من الكتب التعليمية في وصفها على "رخويات أسلاف افتراضية" (انظر الصورة أدناه). تحتوي هذه المجموعة على "مثل-البطلينوس" على قوقعة في الأعلى، وهي مصنوعة من الپروتينات والكايتين معززة بكربونات الكالسيوم، وتُفرزها بواسطة وشاح يغطي السطح العلوي بالكامل. يتكون الجانب السفلي من الحيوان من "قدم" عضلية واحدة.

على الرغم من أن الرخويات من الجوفيات، إلا أن الجوف يميل إلى أن الصغر. تجويف الجسم الرئيسي يكون دورياً الذي من خلاله يدور الدم؛ على هذا النحو، فإن الجهاز الدوري الخاص بالرخويات هو بشكل أساسي مفتوحاً. يتكون الجهاز الهضمي "المعمم" للرخويات من "لسان" خشن، مبرد، وجهاز هضمي معقد يلعب فيه المخاط و"الشعر" المجهري الذي يعمل بالطاقة العضلية والذي يسمى الأهداب أدواراً هامة مختلفة. تحتوي الرخويات المعممة على زوجين من الحبال العصبية، أو ثلاثة في ذوات المصراعين. المخ، في الأنواع التي لديها واحد، يحيط بالمريء.

تحتوي معظم الرخويات على عينين، وجميعها لديها مجسات للكشف عن المواد الكيميائية والاهتزازات واللمس. أبسط نوع من الأجهزة التناسلية للرخويات يعتمد على الإخصاب الخارجي، لكن تحدث اختلافات أكثر تعقيداً. تنتج جميعها تقريباً البيض، والتي قد تخرج منها يرقات حاملة العجل، أو يرقات حاملة الغشاء الأكثر تعقيداً، أو يرقات بالغة مصغرة. يُختزل التجويف الجوفي. لدى الرخويات نظام الدورة الدموية المفتوحة وأعضاء تشبه الكلى للإخراج.

توجد أدلة جيدة على ظهور البطنقدميات، الرأسقدميات، وذوات المصراعين في العصر الكامبري، منذ 541–485.4 مليون سنة. ومع ذلك، فإن التاريخ التطوري لنشوء الرخويات من أسلاف عجلانيات عرفية وتنوعها إلى أشكال حية وأحفورية لا يزال موضع نقاش حاد بين العلماء.

أمونيت متحجرة معروضة في المتحف الوطني الفلپيني.

كانت الرخويات ولا تزال مصدراً غذائياً هاماً للبشر. السموم التي يمكن أن تتراكم في بعض الرخويات في ظل ظروف معينة تخلق خطر التسمم الغذائي، والعديد من الولايات القضائية لديها لوائح للحد من هذا الخطر. كانت الرخويات أيضاً، لعدة قرون، مصدراً للسلع الكمالية الهامة، ولا سيما اللؤلؤ، عرق اللؤلؤ، وصبغة أرجوان صور، وحرير البحر. كما أُستخدمت كنقود في بعض مجتمعات ما قبل الصناعية.

تعتبر حفنة من أنواع الرخويات في بعض الأحيان مخاطر أو آفات للأنشطة البشرية. لدغة الأخطبوط أزرق الحلقات غالباً ما تكون قاتلة، ولدغة أخطبوط المحيط الهادي العملاق تسبب التهاب يمكن أن يستمر لأكثر من شهر. يمكن أن تتسبب لسعات بعض أنواع القواقع المخروطية الإستوائية الكبيرة من فصيلة الحلزونات المخروطية في الوفاة، لكن سمومها المتطورة، على الرغم من سهولة إنتاجها، أصبحت أدوات هامة في الأبحاث العصبية. تنتقل البلهارسيا (المعروفة أيضاً بحمى الحلزون) إلى البشر عن طريق الحلزونات المائية، وتصيب حوالي 200 مليون شخص. يمكن أن تكون القواقع والرخويات أيضاً آفات زراعية خطيرة، وقد أدى الإدخال العرضي أو المتعمد لبعض أنواع القواقع إلى بيئات جديدة إلى إلحاق أضرار جسيمة ببعض الأنظمة البيئية.

أصل الاسم

The words mollusc and mollusk are both derived from the French mollusque, which originated from the post-classical Latin mollusca, from mollis, soft, first used by J. Jonston (Historiæ Naturalis, 1650) to describe a group comprising cephalopods.[10] Molluscus is used in classical Latin as an adjective only with nux (nut) to describe a particular type of soft nut. The use of mollusca in biological taxonomy by Jonston and later Linnaeus may have been influenced by Aristotle's τὰ μαλάκια ta malákia (the soft ones; < μαλακός malakós "soft"), which he applied among other things to cuttlefish.[11][12] The scientific study of molluscs is accordingly called malacology.[13]

The name Molluscoida was formerly used to denote a division of the animal kingdom containing the brachiopods, bryozoans, and tunicates, the members of the three groups having been supposed to somewhat resemble the molluscs. As now known, Brachipoda, Bryozoa and Mollusca are all part of the Lophotrochozoa but have very little relation to the Tunicata, so the name Molluscoida has been abandoned.[14]

التعريف

The most universal features of the body structure of molluscs are a mantle with a significant body cavity used for breathing and excretion, and the organization of the nervous system. Many have a calcareous shell.[15]

Molluscs have developed such a varied range of body structures, finding synapomorphies (defining characteristics) to apply to all modern groups is difficult.[16] The most general characteristic of molluscs is they are unsegmented and bilaterally symmetrical.[17] The following are present in all modern molluscs:[18][20]

Other characteristics that commonly appear in textbooks have significant exceptions:

Whether characteristic is found in these classes of Molluscs
Supposed universal Molluscan characteristic[18] Aplacophora
[19](p. 291–292) 		Polyplacophora
[19](p. 292–298) 		Monoplacophora
[19](p. 298–300) 		Gastropoda
[19](p. 300–343) 		Cephalopoda
[19](p. 343–367) 		Bivalvia
[19](p. 367–403) 		Scaphopoda
[19](p. 403–407)
Radula, a rasping "tongue" with chitinous teeth Absent in 20% of Neomeniomorpha Yes Yes Yes Yes No Internal, cannot extend beyond body
Broad, muscular foot Reduced or absent Yes Yes Yes Modified into arms Yes Small, only at "front" end
Dorsal concentration of internal organs (visceral mass) Not obvious Yes Yes Yes Yes Yes Yes
Large digestive ceca No ceca in some Aplacophora Yes Yes Yes Yes Yes No
Large complex metanephridia ("kidneys") None Yes Yes Yes Yes Yes Small, simple
One or more valves/shells Primitive forms, yes; modern forms, no Yes Yes Snails, yes; slugs, mostly yes (internal vestigial) Octopuses, some (vestigial); cuttlefish, nautilus, squid, yes Yes Yes
Odontophore Yes Yes Yes Yes Yes No Yes

التنوع

Diversity and variability of shells of molluscs on display
About 80% of all known mollusc species are gastropods (snails and slugs), including this cowry (a sea snail).[21]

Estimates of accepted described living species of molluscs vary from 50,000 to a maximum of 120,000 species.[4] The total number of described species is difficult to estimate because of unresolved synonymy. In 1969, David Nicol estimated the probable total number of living mollusc species at 107,000 of which were about 12,000 fresh-water gastropods and 35,000 terrestrial. The Bivalvia would comprise about 14% of the total and the other five classes less than 2% of the living molluscs.[22] In 2009, Chapman estimated the number of described living mollusc species at 85,000.[4] Haszprunar in 2001 estimated about 93,000 named species,[23] which include 23% of all named marine organisms.[24] Molluscs are second only to arthropods in numbers of living animal species[21]—far behind the arthropods' 1,113,000 but well ahead of chordates' 52,000.[19](p. Front endpaper) About 200,000 living species in total are estimated,[4][25] and 70,000 fossil species,[18] although the total number of mollusc species ever to have existed, whether or not preserved, must be many times greater than the number alive today.[26]

Molluscs have more varied forms than any other animal phylum. They include snails, slugs and other gastropods; clams and other bivalves; squids and other cephalopods; and other lesser-known but similarly distinctive subgroups. The majority of species still live in the oceans, from the seashores to the abyssal zone, but some form a significant part of the freshwater fauna and the terrestrial ecosystems. Molluscs are extremely diverse in tropical and temperate regions, but can be found at all latitudes.[16] About 80% of all known mollusc species are gastropods.[21] Cephalopoda such as squid, cuttlefish, and octopuses are among the most neurologically advanced of all invertebrates.[27] The giant squid, which until recently had not been observed alive in its adult form,[28] is one of the largest invertebrates, surpassed in weight but not in length by the colossal squid.[29]

Freshwater and terrestrial molluscs appear exceptionally vulnerable to extinction. Estimates of the numbers of non-marine molluscs vary widely, partly because many regions have not been thoroughly surveyed. There is also a shortage of specialists who can identify all the animals in any one area to species. However, in 2004 the IUCN Red List of Threatened Species included nearly 2,000 endangered non-marine molluscs. For comparison, the great majority of mollusc species are marine, but only 41 of these appeared on the 2004 Red List. About 42% of recorded extinctions since the year 1500 are of molluscs, consisting almost entirely of non-marine species.[30]

التشريح

للمزيد من المعلومات: Mollusc shell and Pollicina
ملف:Archimollusc-en.svg
Anatomical diagram of a hypothetical ancestral mollusc

Because of the great range of anatomical diversity among molluscs, many textbooks start the subject of molluscan anatomy by describing what is called an archi-mollusc, hypothetical generalized mollusc, or hypothetical ancestral mollusc (HAM) to illustrate the most common features found within the phylum. The depiction is visually rather similar to modern monoplacophorans.[16][20][31]

The generalized mollusc is an unsegmented, bilaterally symmetrical animal and has a single, "limpet-like" shell on top. The shell is secreted by a mantle covering the upper surface. The underside consists of a single muscular "foot".[20] The visceral mass, or visceropallium, is the soft, nonmuscular metabolic region of the mollusc. It contains the body organs.[17]

الوشاح وتجويف الوشاح

The mantle cavity, a fold in the mantle, encloses a significant amount of space. It is lined with epidermis, and is exposed, according to habitat, to sea, fresh water or air. The cavity was at the rear in the earliest molluscs, but its position now varies from group to group. The anus, a pair of osphradia (chemical sensors) in the incoming "lane", the hindmost pair of gills and the exit openings of the nephridia (kidneys) known as "Organs of bojanus" and gonads (reproductive organs) are in the mantle cavity.[20] The whole soft body of bivalves lies within an enlarged mantle cavity.[17]

القوقعة

المقالة الرئيسية: قوقعة الرخويات

The mantle edge secretes a shell (secondarily absent in a number of taxonomic groups, such as the nudibranchs[17]) that consists of mainly chitin and conchiolin (a protein hardened with calcium carbonate),[20][32] except the outermost layer, which in almost all cases is all conchiolin (see periostracum).[20] Molluscs never use phosphate to construct their hard parts,[33] with the questionable exception of Cobcrephora.[34] While most mollusc shells are composed mainly of aragonite, those gastropods that lay eggs with a hard shell use calcite (sometimes with traces of aragonite) to construct the eggshells.[35]

The shell consists of three layers: the outer layer (the periostracum) made of organic matter, a middle layer made of columnar calcite, and an inner layer consisting of laminated calcite, often nacreous.[17]

In some forms the shell contains openings. In abalone there are holes in the shell used for respiration and the release of egg and sperm, in the nautilus a string of tissue called the siphuncle goes through all the chambers, and the eight plates that make up the shell of chitons are penetrated with living tissue with nerves and sensory structures.[36] The earliest-derived living mollusca, the Polyplacophora (chitons) and shell-less vermiform Aplacophora, remains contentious despite many developmental and molecular studies of these organisms.[37][38] Both studies investigated molluscan phylogeny through synthesis of paleontological and neontological data,[37] and the other being large and small-subunit nuclear rRna sequences.[38] Another article describing the chitons, with their shells, this time, is one by D K Jacobs and others, where shell evolution of molluscs and chitons are examined to visualize the evolution of the chiton, from the Polyplacophora, to the Aplacophora, and others.[39]

الأقدام

A 50-second video of snails (most likely Natica chemnitzi and Cerithium stercusmuscaram) feeding on the sea floor in the Gulf of California, Puerto Peñasco, Mexico

The body of a mollusc has a ventral muscular foot, which is adapted to different purposes (locomotion, grasping the substratum, burrowing or feeding) in different classes.[40] The foot carries a pair of statocysts, which act as balance sensors. In gastropods, it secretes mucus as a lubricant to aid movement. In forms having only a top shell, such as limpets, the foot acts as a sucker attaching the animal to a hard surface, and the vertical muscles clamp the shell down over it; in other molluscs, the vertical muscles pull the foot and other exposed soft parts into the shell.[20] In bivalves, the foot is adapted for burrowing into the sediment;[40] in cephalopods it is used for jet propulsion,[40] and the tentacles and arms are derived from the foot.[41]

الدورة الدموية

Most molluscs' circulatory systems are mainly open, except for cephalopods, whose circulatory systems are closed. Although molluscs are coelomates, their coeloms are reduced to fairly small spaces enclosing the heart and gonads. The main body cavity is a hemocoel through which blood and coelomic fluid circulate and which encloses most of the other internal organs. These hemocoelic spaces act as an efficient hydrostatic skeleton.[17] The blood of these molluscs contains the respiratory pigment hemocyanin as an oxygen-carrier. The heart consists of one or more pairs of atria (auricles), which receive oxygenated blood from the gills and pump it to the ventricle, which pumps it into the aorta (main artery), which is fairly short and opens into the hemocoel.[20] The atria of the heart also function as part of the excretory system by filtering waste products out of the blood and dumping it into the coelom as urine. A pair of metanephridia ("little kidneys") to the rear of and connected to the coelom extracts any re-usable materials from the urine and dumps additional waste products into it, and then ejects it via tubes that discharge into the mantle cavity.[20]

Exceptions to the above are the molluscs Planorbidae or ram's horn snails, which are air-breathing snails that use iron-based hemoglobin instead of the copper-based hemocyanin to carry oxygen through their blood.

التنفس

Most molluscs have only one pair of gills, or even only a singular gill. Generally, the gills are rather like feathers in shape, although some species have gills with filaments on only one side. They divide the mantle cavity so water enters near the bottom and exits near the top. Their filaments have three kinds of cilia, one of which drives the water current through the mantle cavity, while the other two help to keep the gills clean. If the osphradia detect noxious chemicals or possibly sediment entering the mantle cavity, the gills' cilia may stop beating until the unwelcome intrusions have ceased. Each gill has an incoming blood vessel connected to the hemocoel and an outgoing one to the heart.[20]

التغذية، الهضم، والإخراج

قالب:Annotated image/Snail radula working Molluscs use intracellular digestion. Most molluscs have muscular mouths with radulae, "tongues", bearing many rows of chitinous teeth, which are replaced from the rear as they wear out. The radula primarily functions to scrape bacteria and algae off rocks, and is associated with the odontophore, a cartilaginous supporting organ.[17] The radula is unique to the molluscs and has no equivalent in any other animal.

Molluscs' mouths also contain glands that secrete slimy mucus, to which the food sticks. Beating cilia (tiny "hairs") drive the mucus towards the stomach, so the mucus forms a long string called a "food string".[20]

At the tapered rear end of the stomach and projecting slightly into the hindgut is the prostyle, a backward-pointing cone of feces and mucus, which is rotated by further cilia so it acts as a bobbin, winding the mucus string onto itself. Before the mucus string reaches the prostyle, the acidity of the stomach makes the mucus less sticky and frees particles from it.[20]

The particles are sorted by yet another group of cilia, which send the smaller particles, mainly minerals, to the prostyle so eventually they are excreted, while the larger ones, mainly food, are sent to the stomach's cecum (a pouch with no other exit) to be digested. The sorting process is by no means perfect.[20]

Periodically, circular muscles at the hindgut's entrance pinch off and excrete a piece of the prostyle, preventing the prostyle from growing too large. The anus, in the part of the mantle cavity, is swept by the outgoing "lane" of the current created by the gills. Carnivorous molluscs usually have simpler digestive systems.[20]

As the head has largely disappeared in bivalves, the mouth has been equipped with labial palps (two on each side of the mouth) to collect the detritus from its mucus.[17]

الجهاز العصبي

مخطط مبسط للجهاز العصبي للرخويات.

The cephalic molluscs have two pairs of main nerve cords organized around a number of paired ganglia, the visceral cords serving the internal organs and the pedal ones serving the foot. Most pairs of corresponding ganglia on both sides of the body are linked by commissures (relatively large bundles of nerves). The ganglia above the gut are the cerebral, the pleural, and the visceral, which are located above the esophagus (gullet). The pedal ganglia, which control the foot, are below the esophagus and their commissure and connectives to the cerebral and pleural ganglia surround the esophagus in a circumesophageal nerve ring or nerve collar.[19](p. 284–291)

The acephalic molluscs (i.e., bivalves) also have this ring but it is less obvious and less important. The bivalves have only three pairs of ganglia—cerebral, pedal, and visceral—with the visceral as the largest and most important of the three functioning as the principal center of "thinking".[42][43] Some such as the scallops have eyes around the edges of their shells which connect to a pair of looped nerves and which provide the ability to distinguish between light and shadow.

التكاثر

انظر أيضاً: Reproductive system of gastropods and Reproductive system of cephalopods

قالب:Annotated image/Trochophore larva The simplest molluscan reproductive system relies on external fertilization, but with more complex variations. All produce eggs, from which may emerge trochophore larvae, more complex veliger larvae, or miniature adults. Two gonads sit next to the coelom, a small cavity that surrounds the heart, into which they shed ova or sperm. The nephridia extract the gametes from the coelom and emit them into the mantle cavity. Molluscs that use such a system remain of one sex all their lives and rely on external fertilization. Some molluscs use internal fertilization and/or are hermaphrodites, functioning as both sexes; both of these methods require more complex reproductive systems.[20] C. obtusus is an endemic snail species of the Eastern Alps. There is strong evidence for self-fertilization in the easternmost snail populations of this species.[44]

The most basic molluscan larva is a trochophore, which is planktonic and feeds on floating food particles by using the two bands of cilia around its "equator" to sweep food into the mouth, which uses more cilia to drive them into the stomach, which uses further cilia to expel undigested remains through the anus. New tissue grows in the bands of mesoderm in the interior, so the apical tuft and anus are pushed further apart as the animal grows. The trochophore stage is often succeeded by a veliger stage in which the prototroch, the "equatorial" band of cilia nearest the apical tuft, develops into the velum ("veil"), a pair of cilia-bearing lobes with which the larva swims. Eventually, the larva sinks to the seafloor and metamorphoses into the adult form. While metamorphosis is the usual state in molluscs, the cephalopods differ in exhibiting direct development: the hatchling is a 'miniaturized' form of the adult.[45] The development of molluscs is of particular interest in the field of ocean acidification as environmental stress is recognized to affect the settlement, metamorphosis, and survival of larvae.[46]

علم البيئة

التغذية

Most molluscs are herbivorous, grazing on algae or filter feeders. For those grazing, two feeding strategies are predominant. Some feed on microscopic, filamentous algae, often using their radula as a 'rake' to comb up filaments from the sea floor. Others feed on macroscopic 'plants' such as kelp, rasping the plant surface with its radula. To employ this strategy, the plant has to be large enough for the mollusc to 'sit' on, so smaller macroscopic plants are not as often eaten as their larger counterparts.[47] Filter feeders are molluscs that feed by straining suspended matter and food particles from water, typically by passing the water over their gills. Most bivalves are filter feeders, which can be measured through clearance rates. Research has demonstrated that environmental stress can affect the feeding of bivalves by altering the energy budget of organisms.[46]

Cephalopods are primarily predatory, and the radula takes a secondary role to the jaws and tentacles in food acquisition. The monoplacophoran Neopilina uses its radula in the usual fashion, but its diet includes protists such as the xenophyophore Stannophyllum.[48] Sacoglossan sea-slugs suck the sap from algae, using their one-row radula to pierce the cell walls,[49] whereas dorid nudibranchs and some Vetigastropoda feed on sponges[50][51] and others feed on hydroids.[52] (An extensive list of molluscs with unusual feeding habits is available in the appendix of قالب:Molluscan diets)

التصنيف

انظر أيضاً: قائمة رتب الرخويات

Opinions vary about the number of classes of molluscs; for example, the table below shows seven living classes,[23] and two extinct ones. Although they are unlikely to form a clade, some older works combine the Caudofoveata and Solenogastres into one class, the Aplacophora.[31][19](p. 291–292) Two of the commonly recognized "classes" are known only from fossils.[21]

Class Major organisms Described living species[23] Distribution
Gastropoda [19](p. 300) all snails and slugs including abalone, limpets, conch, nudibranchs, sea hares, sea butterflies 70,000 marine, freshwater, land
Bivalvia [19](p. 367) clams, oysters, scallops, geoducks, mussels, rudists† 20,000 marine, freshwater
Polyplacophora [19](pp. 292–8) chitons 1,000 rocky tidal zone and seabed
Cephalopoda [19](p. 343) squid, octopuses, cuttlefish, nautiluses, vampire squids, Spirula, belemnites†, ammonites† 900 marine
Scaphopoda [19](pp. 403–7) tusk shells 500 marine 6–7،000 متر (20–22،966 ft)
Aplacophora [19](pp. 291–2) worm-like molluscs 320 seabed 200–3،000 متر (660–9،840 ft)
Monoplacophora [19](pp. 298–300) ancient lineage of molluscs with cap-like shells 31 seabed 1،800–7،000 متر (5،900–23،000 ft); one species 200 متر (660 ft)
Rostroconchia†[53] fossils; probable ancestors of bivalves extinct marine
Helcionelloida†[54] fossils; snail-like molluscs such as Latouchella extinct marine

الشجرة التطورية

قالب:Thumb The phylogeny (evolutionary "family tree") of molluscs is a controversial subject. In addition to the debates about whether Kimberella and any of the "halwaxiids" were molluscs or closely related to molluscs,[55][56][57][58] debates arise about the relationships between the classes of living molluscs.[37] In fact, some groups traditionally classified as molluscs may have to be redefined as distinct but related.[59]

Molluscs are generally regarded as members of the Lophotrochozoa,[37] a group defined by having trochophore larvae and, in the case of living Lophophorata, a feeding structure called a lophophore. The other members of the Lophotrochozoa are the annelid worms and seven marine phyla.[60] The diagram on the right summarizes a phylogeny presented in 2007 without the annelid worms.

Because the relationships between the members of the family tree are uncertain, it is difficult to identify the features inherited from the last common ancestor of all molluscs.[61] For example, it is uncertain whether the ancestral mollusc was metameric (composed of repeating units)—if it was, that would suggest an origin from an annelid-like worm.[39] Scientists disagree about this: Giribet and colleagues concluded, in 2006, the repetition of gills and of the foot's retractor muscles were later developments,[16] while in 2007, Sigwart concluded the ancestral mollusc was metameric, and it had a foot used for creeping and a "shell" that was mineralized.[37] In one particular branch of the family tree, the shell of conchiferans is thought to have evolved from the spicules (small spines) of aplacophorans; but this is difficult to reconcile with the embryological origins of spicules.[61]

The molluscan shell appears to have originated from a mucus coating, which eventually stiffened into a cuticle. This would have been impermeable and thus forced the development of more sophisticated respiratory apparatus in the form of gills.[54] Eventually, the cuticle would have become mineralized,[54]using the same genetic machinery (engrailed) as most other bilaterian skeletons.[39] The first mollusc shell almost certainly was reinforced with the mineral aragonite.[32]

Classification into higher taxa for molluscan classes has been and remains problematic. Numerous different clades have been proposed but few have received strong support. Traditionally, Mollusca is split into two subphyla, Conchifera and Aculifera, based on the presence of a shell. The "Testaria" hypothesis is similar, but includes chitons with the rest of the conchiferans. Some studies completely reject the proposal, instead favoring a "Serialia" hypothesis which classifies chitons and monoplacophorans as closely related.

Morphological analyses tend to recover a conchiferan clade that receives less support from molecular analyses,[62] although these results also lead to unexpected paraphylies, for instance scattering the bivalves throughout all other mollusc groups.[38]

However, an analysis in 2009 using both morphological and molecular phylogenetics comparisons concluded the molluscs are not monophyletic; in particular, Scaphopoda and Bivalvia are both separate, monophyletic lineages unrelated to the remaining molluscan classes; the traditional phylum Mollusca is polyphyletic, and it can only be made monophyletic if scaphopods and bivalves are excluded.[59] A 2010 analysis recovered the traditional conchiferan and aculiferan groups, and showed molluscs were monophyletic, demonstrating that available data for solenogastres was contaminated.[63] Current molecular data are insufficient to constrain the molluscan phylogeny, and since the methods used to determine the confidence in clades are prone to overestimation, it is risky to place too much emphasis even on the areas of which different studies agree.[64] Rather than eliminating unlikely relationships, the latest studies add new permutations of internal molluscan relationships, even bringing the conchiferan hypothesis into question.[65]


صفوف الرخويات

تضمن الرخويات مجموعة الصفوف التالية:

صف أحادية المصراع أو البطن قدميات

البطن قدميات Gastropoda، هي أكبر صفوف الرخويات وتشمل البطلينوسات، والحلزونات والقواقع والوَلَكْ. ومعظم أنواع أحادية المصراع مزودة بصدفة حلزونية واحدة. ولكن بعض أنواع أحادية المصراع ـ مثل حلزونات الحديقة وحلزونات البحر المسماة عارية الخياشيم ـ ليس لديها صدفة بعد الطور اليرقي. وتوجد فصيلة مهمة من أحاديات المصراع في أستراليا، تسمى فصيلة الهيليكارونيدي، تمثل مرحلة انتقالية بين الحلزونات والقواقع. ويظهر بعضها مزودًا بصدفة مختزلة جدًا ولامعة.[66]

تبدو بطنية الأقدام وكأنها تزحف على بطونها ولكنها في الحقيقة تستخدم قدمًا عضليًا كبيرًا. ويمتد هذا القدم تحت الجسم وتتحرك عضلاته بطريقة تَمَوجية تجعل الحيوان يتحرك للأمام. وتملك معظم القواقع البحرية وبعض القواقع الأرضية تركيبًا يشبه الغطاء على ظهر القدم يسمى غطاء الصدفة. وعندما يُهدد خطرُ ما القوقع، فإنه ينسحب للخلف داخل الصدفة ويُغلق غطاء الصدفة.

وبعض أنواع أحادية المصراع لها زوجان من اللوامس (المجسات) على رأسها. ويساعد أحد الزوجين الحيوان في تحسس طريقه. ويحمل كل من لامسي الزوج الآخر عينًا في بعض الأنواع، وهناك أنواع أخرى لا تملك عيونًا على الإطلاق. وكل حيوان أحادي المصراع مزود أيضًا بشريط من الأسنان يسمى الزائدة الكاشطة يعمل كمبرد خشن يكشط غذاء الحيوان.

ومعظم أحاديات المصراع، التي تتغذى بالنباتات، مزودة بالآلاف من الأسنان الضعيفة. وثمة أنواع قليلة، تتغذى بالرخويات الأخرى، بها عدة رُزم من الأسنان القوية.

صف ثنائية المصراع (ذوات المصراعين، بُليطية الأقدام)

ثنائيات المصراع Bivalvia تُكوِن ثانية كبرى مجموعات الرخويات وتشمل المحار الملزمي والمحار وبلح البحر والأسقالوب (المحار المروحي) و ديدان السُفُن. وكل ذوات المصراعين مزودة بصدفتين متصلتين وعندما يُفَزَّع الحيوان تقوم عضلات قوية بجذب الصدفتين لتقفلهما وتبقيهما مقفلتين حتى زوال الخطر. ومعظم ذوات المصراعين صغيرة الحجم ولكن بعضها كبير الحجم، مثل المحار الملزمي العملاق الذي قد ينمو لأكثر من متر طولاً ويزن حوالي 1,100 كجم.

ولدى ذوات المصراعين قدم عضلية قوية وتتحرك أنواع كثيرة بدفع القدم للخارج وتثبيته في الطين أو الرمل، وبعد ذلك تقتلع نفسها مع القدم. وبعض ذوات المصراعين ،مثل بطة الأرض والمحار الملزمي الشفري تستعمل القدم في حفر الفجوات. فهي تدفع القدم لأسفل في الطين أو الرمل. وفي البداية تتمدد القدم لتوسع الحفرة، ثم تنقبض بعد ذلك لتجذب الصدفة داخل الحفرة. ويستطيع محار الفولاس أن يحفر فجوات حتى في الطين الجامد أو الصخور الناعمة.

وذوات المصراعين ليس لديها رأس أو أسنان. وهي تحصل على الأكسجين والطعام عن طريق أنبوب عضلي (سيفون). ويستطيع الأنبوب أن يمتد ليصل إلى الطعام والماء إذا كان الحيوان مدفونًا في الطين أو الرمل. وتتغذى ذوات المصراعين بالمواد النباتية التي تُرشَّح من الماء بوساطة الخياشيم.

صف الأخطبوطات والحبَّار (رأسيات الأقدام)

رأسقدميات Cephalopoda، هي أنشط الرخويات وتشمل الأرغونوط والصَّبيِّد والنوتي. وتعيش كل هذه الرخويات في المحيطات.

يبدو الحيوان رأسي القدم وكأنه مكوَّن من رأس كبير وأذرع طويلة تبدو مثل الأقدام. ويملك الأخطبوط والحبَّار رأسًا قُبيِّة الشكل محاطة بالأذرع. والأخطبوط له ثماني أذرع أما الحبَّار فله ثماني أذرع ولامسان. وتنمو الأذرع حول فكين صلبين قويين متقاريي الشكل يوجدان أسفل الرأس. ويقوم هذان الفكان بتمزيق فريسة الحيوان، وهما أشد خطورة من الأذرع. وتستعمل الأخطبوطات أذرعها بينما يستعمل الحبار كل لوامسه وأذرعه في القبض على الفريسة وجذبها عبر فكيه. ويتغذى الأخطبوط والحبَّار بالأسماك والرخويات الأخرى والأسماك الصدفية.

صف الأصداف السِّنْيّة (زورقية الأقدام)

زورقية الأقدام Scaphopoda، لها أصداف رفيعة ملتوية تشبه أسنان الفيل وغالبًا ما تسمى هذه الرخويات بأصداف أسنان الفيل. ولدى حيوان الصدفة السِّنية قدم مديدة تشبه إلى حد ما زورقًا صغيرًا.

تعيش كل الحيوانات ذات الأصداف السِّنيِّة في المحيطات، حيث تحفر في الطين أو الرمل في قاع المحيط، بينما تبرز قمة الصدفة في الماء. وليس لدى الحيوانات ذات الأصداف السنية رؤوس ولا عيون، وتتغذى بالحيوانات والنباتات ذات الخلية الواحدة التي تدفعها داخل فمها بواسطة اللوامس.

صف الخيتونات أو عديدات الأصداف

عديدات الأصداف Chiton، حيوان بحري ذو درع واقية صلبة. تتكون دروع معظمالخَيْتُونَات من ثماني قطع متداخلة، يربطها ببعضها لحم كالجلد. ويوجد أكثر من 800 نوع من الخَيْتُونَات اليوم. وهي تعيش في المياه الضحلة في معظم أنحاء العالم. يقلّ طول بعض أنواع الخَيْتُون عن سنتيمترين ونصف السنتيمتر، بينما تطول أنواع أخرى مثل خَيْتُون ستيلير، حتى تبلغ 30سم. للخَيْتُون عضو عضلي كبير ومفلطح، يسمى قدمًا، يستخدمه للالتصاق بالصخور عن طريق الامتصاص. تقتات بعض الخيتونات الأحياء النباتية فقط، وبعضها الآخر يقتات الحيوانات البحرية الصغيرة. [67]

وهي ذات أجسام بيضية مفلطحة مغطاة بثماني صفائح صدفية مثبتة بعضها مع بعض بحزام متين. ويشير اسم عديدة الأصداف إلى صدفة الخيتون المكونة من ثماني قطع متداخلة مع بعضها. ولدى الخيتون قدم كبيرة مسطحة يستخدمها في الحركة، ولكنه عادة ما يلتصق بالصخور بإحكام، وحينما يجبر على ترك صخرته، فإنه يتكور في شكل كرة. ويملك الخيتون رأسًا صغيرًا وفمًا، ولكن ليس له عيون أو لوامس. ويستعمل الخيتون زائدته الكاشطة الطويلة ذات الأسنان العديدة، لكشط الأعشاب البرية من الصخور للغذاء.

صف أحاديات الصدفة

أحاديات الصدفة Monoplacophora، توجد معظمها في شكل متحجِّرات فقط. وأحاديات الصدفة حيوانات ذات صدفة واحدة مسطحة تقريبًا شبيهة بصدفة البطلينوس. وهي شاذة عن بقية الرخويات في أن لها العديد من أزواج الخياشيم، وفي أن لها ستة أزواج أو أكثر من الكلى ولها جهاز عصبي مركزي شبيه بالسُلَّم. ولكن لها مثل بقية الرخويات بُرْنسًا، وزائدة كاشطة. ولا يعرف الكثير عن عاداتها.

صف عديمات الأصداف

عديمات الأصداف Aplacophora، وهي رخويات دودية الشكل مغطاة بأشواك صغيرة وهي نادرًا ماترى.

التاريخ التطوري

المقالة الرئيسية: تطور الرخويات
انظر أيضاً: Evolution of cephalopods
The enigmatic Kimberella quadrata (fossil pictured) from the Ediacaran has been described as being "mollusc-like" because of its features which are shared with modern day molluscs.

Good evidence exists for the appearance of gastropods (e.g., Aldanella), cephalopods (e.g., Plectronoceras) and bivalves (Pojetaia, Fordilla) towards the middle of the Cambrian period, c. 500 مليون سنة مضت, though arguably each of these may belong only to the stem lineage of their respective classes.[68] However, the evolutionary history both of the emergence of molluscs from the ancestral group Lophotrochozoa, and of their diversification into the well-known living and fossil forms, is still vigorously debated.

Debate occurs about whether some Ediacaran and Early Cambrian fossils really are molluscs.[69] Kimberella, from about 555 مليون سنة مضت, has been described by some paleontologists as "mollusc-like",[70][55] but others are unwilling to go further than "probable bilaterian",[56][37] if that.[71]

There is an even sharper debate about whether Wiwaxia, from about 505 مليون سنة مضت, was a mollusc, and much of this centers on whether its feeding apparatus was a type of radula or more similar to that of some polychaete worms.[56][57] Nicholas Butterfield, who opposes the idea that Wiwaxia was a mollusc, has written that earlier microfossils from 515 to 510 مليون سنة مضت are fragments of a genuinely mollusc-like radula.[58] This appears to contradict the concept that the ancestral molluscan radula was mineralized.[72]

The tiny helcionellid fossil Yochelcionella is thought to be an early mollusc.[54]
Spirally coiled shells appear in many gastropods.[19](pp. 300–343)

However, the helcionellids, which first appear over 540 مليون سنة مضت in Early Cambrian rocks from Siberia and China,[73][74] are thought to be early molluscs with rather snail-like shells. Shelled molluscs therefore predate the earliest trilobites.[54] Although most helcionellid fossils are only a few millimeters long, specimens a few centimeters long have also been found, most with more limpet-like shapes. The tiny specimens have been suggested to be juveniles and the larger ones adults.[75]

Some analyses of helcionellids concluded these were the earliest gastropods.[76] However, other scientists are not convinced these Early Cambrian fossils show clear signs of the torsion that identifies modern gastropods twists the internal organs so the anus lies above the head.[19](pp. 300–343)[77][78]

  = Septa
  = Siphuncle
كبـِّر
Septa and siphuncle in nautiloid shell

Volborthella, some fossils of which predate 530 مليون سنة مضت, was long thought to be a cephalopod, but discoveries of more detailed fossils showed its shell was not secreted, but built from grains of the mineral silicon dioxide (silica), and it was not divided into a series of compartments by septa as those of fossil shelled cephalopods and the living Nautilus are. Volborthella's classification is uncertain.[79] The Middle Cambrian fossil Nectocaris was interpreted as a cephalopod with 2 arms and no shell by some researchers, but it is later reinterpreted as relative of modern chaetognaths (arrow worms).[80] The Late Cambrian fossil Plectronoceras is now thought to be the earliest undisputed cephalopod fossil, as its shell had septa and a siphuncle, a strand of tissue that Nautilus uses to remove water from compartments it has vacated as it grows, and which is also visible in fossil ammonite shells. However, Plectronoceras and other early cephalopods crept along the seafloor instead of swimming, as their shells contained a "ballast" of stony deposits on what is thought to be the underside, and had stripes and blotches on what is thought to be the upper surface.[81] All cephalopods with external shells except the nautiloids became extinct by the end of the Cretaceous period 65 مليون سنة مضت.[82] However, the shell-less Coleoidea (squid, octopus, cuttlefish) are abundant today.[83]

The Early Cambrian fossils Fordilla and Pojetaia are regarded as bivalves.[84][85][86][87] "Modern-looking" bivalves appeared in the Ordovician period, 488 to 443 مليون سنة مضت.[88] One bivalve group, the rudists, became major reef-builders in the Cretaceous, but became extinct in the Cretaceous–Paleogene extinction event.[89] Even so, bivalves remain abundant and diverse.

The Hyolitha are a class of extinct animals with a shell and operculum that may be molluscs. Authors who suggest they deserve their own phylum do not comment on the position of this phylum in the tree of life.[90]

التفاعل مع البشر

المقالة الرئيسية: الرخويات في الثقافة

For millennia, molluscs have been a source of food for humans, as well as important luxury goods, notably pearls, mother of pearl, Tyrian purple dye, sea silk, and chemical compounds. Their shells have also been used as a form of currency in some preindustrial societies. Some species of molluscs can bite or sting humans, and some have become agricultural pests.

الاستخدام البشري

للمزيد من المعلومات: قوقعة بحرية and قائمة الرخويات الصالحة للأكل

Molluscs, especially bivalves such as clams and mussels, have been an important food source since at least the advent of anatomically modern humans, and this has often resulted in overfishing.[91] Other commonly eaten molluscs include octopuses and squids, whelks, oysters, and scallops.[92] In 2005, China accounted for 80% of the global mollusc catch, netting almost 11،000،000 طن متري (11،000،000 long ton; 12،000،000 short ton). Within Europe, France remained the industry leader.[93] Some countries regulate importation and handling of molluscs and other seafood, mainly to minimize the poison risk from toxins that can sometimes accumulate in the animals.[94]

Photo of three circular metal cages in shallows, with docks, boathouses and palm trees in background
Saltwater pearl oyster farm in Seram, Indonesia

Most molluscs with shells can produce pearls, but only the pearls of bivalves and some gastropods, whose shells are lined with nacre, are valuable.[19](pp. 300–343, 367–403) The best natural pearls are produced by marine pearl oysters, Pinctada margaritifera and Pinctada mertensi, which live in the tropical and subtropical waters of the Pacific Ocean. Natural pearls form when a small foreign object gets stuck between the mantle and shell.

The two methods of culturing pearls insert either "seeds" or beads into oysters. The "seed" method uses grains of ground shell from freshwater mussels, and overharvesting for this purpose has endangered several freshwater mussel species in the southeastern United States.[19](pp. 367–403) The pearl industry is so important in some areas, significant sums of money are spent on monitoring the health of farmed molluscs.[95]

Mosaic of mustachioed, curly-haired man wearing crown and surrounded by halo
Byzantine Emperor Justinian I clad in Tyrian purple and wearing numerous pearls

Other luxury and high-status products were made from molluscs. Tyrian purple, made from the ink glands of murex shells, "fetched its weight in silver" in the fourth century BC, according to Theopompus.[96] The discovery of large numbers of Murex shells on Crete suggests the Minoans may have pioneered the extraction of "imperial purple" during the Middle Minoan period in the 20th–18th centuries BC, centuries before the Tyrians.[97][98] Sea silk is a fine, rare, and valuable fabric produced from the long silky threads (byssus) secreted by several bivalve molluscs, particularly Pinna nobilis, to attach themselves to the sea bed.[99] Procopius, writing on the Persian wars circa 550 CE, "stated that the five hereditary satraps (governors) of Armenia who received their insignia from the Roman Emperor were given chlamys (or cloaks) made from lana pinna. Apparently, only the ruling classes were allowed to wear these chlamys."[100]

Mollusc shells, including those of cowries, were used as a kind of money (shell money) in several preindustrial societies. However, these "currencies" generally differed in important ways from the standardized government-backed and -controlled money familiar to industrial societies. Some shell "currencies" were not used for commercial transactions, but mainly as social status displays at important occasions, such as weddings.[101] When used for commercial transactions, they functioned as a means of exchange similar to money in ordinary business transactions, a tradable good whose value differed from place to place, often as a result of difficulties in transport, and which was vulnerable to incurable inflation if more efficient transport or "goldrush" behavior appeared.[102]

المؤشرات الحيوية

Bivalve molluscs are used as bioindicators to monitor the health of aquatic environments in both fresh water and the marine environments. Their population status or structure, physiology, behaviour or the level of contamination with elements or compounds can indicate the state of contamination status of the ecosystem. They are particularly useful since they are sessile so that they are representative of the environment where they are sampled or placed.[103] Potamopyrgus antipodarum is used by some water treatment plants to test for estrogen-mimicking pollutants from industrial agriculture. Several species of mollusca have been used as bioindicators of environmental stresses that can cause DNA damage. These species include the American oyster Crassostrea virginica,[104] zebra mussels (Dreissena polymorpha)[105][106] and the blue mussel Mytilus edulis.[107]

الإضرار بالبشر

اللدغات والعضات

The blue-ringed octopus's rings are a warning signal; this octopus is alarmed, and its bite can kill.[108]

Some molluscs sting or bite, but deaths from mollusc venoms total less than 10% of those from jellyfish stings.[109]

All octopuses are venomous,[110] but only a few species pose a significant threat to humans. Blue-ringed octopuses in the genus Hapalochlaena, which live around Australia and New Guinea, bite humans only if severely provoked,[108] but their venom kills 25% of human victims. Another tropical species, Octopus apollyon, causes severe inflammation that can last for over a month even if treated correctly,[111] and the bite of Octopus rubescens can cause necrosis that lasts longer than one month if untreated, and headaches and weakness persisting for up to a week even if treated.[112]

Photo of cone on ocean bottom
Live cone snails can be dangerous to shell collectors, but are useful to neurology researchers.[113]

All species of cone snails are venomous and can sting painfully when handled, although many species are too small to pose much of a risk to humans, and only a few fatalities have been reliably reported. Their venom is a complex mixture of toxins, some fast-acting and others slower but deadlier.[113][109][114] The effects of individual cone-shell toxins on victims' nervous systems are so precise as to be useful tools for research in neurology, and the small size of their molecules makes it easy to synthesize them.[113][115]

كناقلات للأمراض

Skin vesicles created by the penetration of Schistosoma (Source: CDC)

Schistosomiasis (also known as bilharzia, bilharziosis or snail fever), a disease caused by the fluke worm Schistosoma, is "second only to malaria as the most devastating parasitic disease in tropical countries. An estimated 200 million people in 74 countries are infected with the disease—100 million in Africa alone."[116] The parasite has 13 known species, two of which infect humans. The parasite itself is not a mollusc, but all the species have freshwater snails as intermediate hosts.[117]

الآفات

Some species of molluscs, particularly certain snails and slugs, can be serious crop pests,[118] and when introduced into new environments, can unbalance local ecosystems. One such pest, the giant African snail Achatina fulica, has been introduced to many parts of Asia, as well as to many islands in the Indian Ocean and Pacific Ocean. In the 1990s, this species reached the West Indies. Attempts to control it by introducing the predatory snail Euglandina rosea proved disastrous, as the predator ignored Achatina fulica and went on to extirpate several native snail species instead.[119]

انظر أيضاً

الهوامش

  1. ^ Most members of this clade are often linked to aculiferans
  2. ^ The formerly dominant British spelling mollusk is still used in the United States — see the reasons given by Gary Rosenberg (1996).[5] For the spelling mollusc, see the reasons given in: Brusca & Brusca. Invertebrates (2nd ed.).

المراجع

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