اسفنج دریایی

از ویکی‌پدیا، دانشنامهٔ آزاد
پرش به ناوبری پرش به جستجو
اسفنج دریایی
محدودهٔ زمانی: ادیاکاران - امروزه
حوزه: یوکاریوت‌ها
فرمانرو: جانوران
شاخه: اسفنج‌تباران
گرنت در تاد، ۱۸۳۶

اسفنج‌های آهکی
اسفنج‌های شیشه‌ای
اسفنج‌های شاخی

اِسفَنج‌های دریایی از جانوران آبزی هستند. سه ویژگی مهم اسفنج‌ها اینست که وسیله حرکتی ندارند، ناجنبنده‌اند و شکل معینی ندارند.

اسفنج‌های دریایی از نظر علمی شاخه‌ای از جانوران را تشکیل می‌دهند که اسفنج‌تباران (Spongiaria) یا روزنک‌تباران (Porifera) نامیده می‌شود. این شاخه، شاخه‌ای از جانوران آبزی غالباً دریازی پالیده‌خوار است که استخوان‌بندی آن‌ها از رشته‌های کلاژنی و سوزنه‌های (spicules) کلسیمی یا سیلیسی تشکیل شده‌است و یاخته‌های تخصصی آن‌ها به‌صورت بافت سازماندهی نشده‌است.[۱]

اسفنج‌ها، موجوداتی بی‌مهره هستند که در دریا زندگی می‌کنند. تا قرن گذشته بعضی از مردم اسفنج‌ها را به اشتباه جزو گیاهان می‌دانستند، امّا در اصل اسفنج غذاسازی نمی‌کند و بنابراین اسفنج‌ها در دستهٔ جانوران محسوب می‌شوند. این جانوران از هر لحاظ ساده هستند. اسفنج‌ها به‌طور کلّی توانایی حرکت ندارند امّا در نمونه‌های نادر توانایی خزیدن دیده می‌شود. این جانوران نه اندامی برای بینایی دارند و نه اندامی برای شنوایی امّا باز هم در نمونه‌های نادر توانایی عکس‌العمل در برابر نور و روشنایی را دارند.

اسفنج‌ها پریاختگانی هستند که به حالت ثابت زیست نموده و دارای اسکلت داخلی کلسیتی یا سیلیسی می‌باشند. پدیده‌های جانشینی مانند سیلیسی شدن و کلسیتی شدن تا حدودی تشخیص انواع فسیل‌های آهکی اسفنج‌ها را از اسفنج‌های سیلیسی مشکل می‌نماید.

به‌طور کلی اسفنج‌ها در دریاها (آب‌های شور) یا آب‌های شیرین زندگی می‌کنند. در کف دریاهای کنونی توده‌های عظیمی از اسفنج‌ها پراکنده هستند. انواع مختلفی از اسفنج‌های آهکی در تمام مناطق کره زمین زندگی می‌کنند. اسفنج‌ها چون ار جانوران ثابت هستند به اجسام سخت احتیاج دارند؛ بنابراین تجمع آن‌ها در حوضچه‌هایی که کف آن از قلوه‌سنگ، تخته سنگ، ریگ و شن و ماسه پوشیده شده باشد بیشتر است.

اسفنج‌ها در آب‌های نیم‌کرهٔ جنوبی گسترش بیشتری دارند؛ و در حقیقت در تمام آب‌های نیم‌کرهٔ جنوبی به عمق ۱۰۰ تا ۵۰۰ متری سکونت دارند و این موضوع به علت حرکت کوه‌های یخی است که به‌طور دائم به اقیانوس وارد می‌شوند و شرایط لازم را برای زیستن اسفنج‌ها فراهم می‌سازند. اسفنج‌های آهکی در اعماق کم و اسفنج‌های سیلیسی در اعماق زیاد زندگی می‌کنند. اسفنج‌های دارای سوزنه‌های (اسپیکول‌های) چهارمحوری و شاخی سیلیسی هم در اعماق کم و هم در اعماق زیاد زندگی می‌کنند. بررسی‌های کنونی نشان داده‌است که بعضی از اسفنج‌ها تا عمق ۶۰۰۰ متری دریا می‌توانند زندگی کنند. این قبیل اسفنج‌ها دارای تیغه‌های درازی نظیر خارداران هستند و شکل آن‌ها نیز کشیده می‌باشد. اصولاً در آب‌های سرد مثل قطب‌ها اسفنج‌ها پراکندگی بیشتری دارند. اسفنج‌های شاخی اصولاً در آب‌های استوایی تا عرض جغرافیایی ۴۵ درجه پراکندگی دارند. از اسفنج‌های شاخی و اسپونژینی اثری خارج از این حوزه نیست. در اسفنج‌های دیگر، هر قدر به طرف قطبین نزدیک شویم سوزنه‌ها درشت‌تر می‌شوند. شوری آب تأثیر زیادی در زندگی اسفنج‌ها دارد. اگر میزان شوری تغییر یابد، تعداد آن‌ها نیز کاسته خواهد شد. جریان آب نیز تأثیر زیادی در زندگی اسفنج‌ها دارد. تکثیر اسفنج‌ها در مناطقی که آب جریان دارد بیشتر است.

شکل اسفنج‌ها نسبت به محیط زیست‌شان متغیر است. شکل اسفنج‌های داخل خلیج‌ها بادبزنی با صفحات چین‌خورده، در آب‌های متلاطم به شکل انگشتی و اسفنج‌های مناطق کم‌عمق، پوستی یا بالشی می‌باشند. در زمان جزر و مد دریاها موقعی‌که اسفنج‌ها در خشکی قرارگیرند، دهان‌هایشان را می‌بندند تا از خروج آب از بدن‌شان جلوگیری شود. به‌طور کلی اسفنج‌های امروزی در منطقه کرانه‌ای تا نسبتاً عمیق دریا زندگی می‌کنند و هیچ‌وقت کلنی حقیقی را تشکیل نمی‌دهند. مطالعهٔ اسفنج‌ها در گذشته نشان می‌دهد که زندگی اسفنج‌ها در گذشته از نظر عمق‌سنجی با انواع کنونی فرق می‌کند. مثلاً در سنگ‌های دوره کرتاسه غالباً در رخساره‌های ساحلی پیدا می‌شوند ولی در زمان‌های بعدی به‌تدریج به اعماق زیادتر مهاجرت نموده‌اند؛ بنابراین می‌توان نتیجه گرفت که اسفنج‌ها در گذشته از اعماق کم به اعماق زیادتر مهاجرت داشته‌اند. از طرفی اسفنج‌های امروزی غالباً انفرادی هستند ولی در انواع گذشته زندگی آن‌ها به حالت کلنی و آبسنگی بوده‌است و سنگواره‌های آن‌ها با آبسنگ‌های مرجانی و خزه‌زی و غیره همراه می‌باشند. در علم دیرینه‌شناسی از فسیل سوزن اسفنج‌ها در تشخیص نوع محیط رسوبی استفاده می‌کنند. اسفنج‌ها ساده‌ترین حیوانات پر سلولی هستند و به عبارت دیگر پست‌ترین یا ابتدایی‌ترین جانوران پر سلولی‌اند. به‌طور کلی محیط زیست آن‌ها آبی و غالباً دریایی بودند و به جسم خارجی متصل‌اند. بدن اسفنج‌ها شبیه یک کیسه تو خالی است که در قاعده به وسیله یک پایه یا بدون پایه به جسمی متصل شده‌است. دیواره آن پر از سوراخ‌های ریز است و از این جهت به آن‌ها روزنک‌تباران می‌گویند.

اسفنج‌ها هم از راه جوانه زدن و هم از راه جنسی تولید مثل می‌کنند.[۲]


اسکلت خارجی این جانوران به‌طور معمول از جنس آهک است. امّا رنگ‌های اسفنج‌ها رنگ‌های گوناگونی دارند و در یک یا دو رنگ منحصر نمی‌شوند. البتّه این رنگ‌ها بستگی به زیستگاهشان هم دارد. مثلاً اسفنج دریاهای مناطق گرمسیر رنگ‌های بسیار درخشانی دارند.

اسفنج‌ها ساده‌ترین حیوانات پرسلولی هستند و به عبارت دیگر پست‌ترین یا ابتدایی‌ترین جانوران پر سلولی‌اند. به‌طور کلی محیط زیست آن‌ها آبی و غالباً دریایی بودند و به جسم خارجی متصل‌اند. بدن اسفنج‌ها شبیه یک کیسه تو خالی است که در قاعده به وسیله یک پایه یا بدون پایه به جسمی متصل شده‌است. دیواره آن پر از سوراخ‌های ریز است و از این جهت به آن‌ها روزنک‌تباران گفته می‌شود. حفره میانی را اسپونگوسل گویند که خود از بالا توسط منفذی به نام اسکولوم (دهانه) به خارج باز می‌شود. اسپونوگوسل در برخی از اسفنج‌ها بزرگ و عمیق و در بعضی دیگر کوچک و کم عمیق است. از نظر شکل بسیار متفاوتند و ممکن است به اشکال مختلف، استوانه‌ای، منشوری، کروی، مخروطی، گلدانی شکل، دیسکی شکل و… دیده شوند. برخی از آن‌ها دارای پایه کوتاه یا بلند هستند. شکل حیوان ن در برخی ثابت است و در عده‌ای دیگر ممکن است متغیر باشد. ابعاد آن غالباً بین ۱ میلی‌متر تا ۲ متر در قطر و ارتفاع تغییر می‌کند. اسفنج‌های عهد حاضر اکثراً به رنگ خاکستری یا خرمایی کم رنگ دیده می‌شوند اما گاهی به رنگ‌های قرمز روشن، آبی، بنفش و سیاه نیز دیده شده‌اند.

نوع تغذیه[ویرایش]

اسفنج‌ها، دستگاهی برای گوارش غذا وبه‌طور کلّی دستگاهی در بدن ندارند. بر روی پوست سخت بدن اسفنج سوراخ‌هایی وجود دارد که به طریق آن‌ها آب به داخل اسفنج وارد می‌شود. این سوراخ‌ها را سلّول‌هایی به نام اسپیکول ساخته‌است. درون سوراخ‌ها را سلّول‌هایی به نام یقه‌دار فرش کرده‌اند. این سلّول‌ها باعث جریان آب در سوراخ‌ها می‌شوند. آبی که به وسیله این سوراخ‌ها و جریان‌ها وارد بدن اسفنج می‌شود، حاوی مواد ریز غذایی و اکسیژن محلول درآب دریا است. وقتی سلّول‌های یقه‌دار غذا را جذب خود کردند، آن را به سلّول‌های آمیب شکل که سرگردان هستند تحویل می‌دهند. این سلّول‌های آمیبی شکل همان‌طور که گفتیم سرگردان هستند. وقتی مواد غذایی ریز را دریافت کردند به نقاطی از بدن اسفنج که نیاز به غذا دارند می‌رساند. مواد زائد این آب که استفاده غذایی ندارند، از سوراخ بزرگ بالای اسفنج خارج می‌شوند. این سوراخ بزرگ اسکولوم نام دارد.

نکاتی جالب دربارهٔ اسفنج‌ها[ویرایش]

  • اکثر مردم اسفنج و مرجان را با هم اشتباه می‌گیرند. تفاوت عمده اسفنج با مرجان نرم بودن اسفنج و سفت و سخت بودن مرجان است.
  • از اسفنج برای مصارف بهداشتی مثل لیف حمّام هم استفاده می‌شود. برای لیف حمّام اوّل صیّادان اسفنج را گرفته، سپس آن را آن قدر لِه می‌کنند تا همه سلّول‌هایش بمیرند تا فقط اسکلت خارجی آن بماند.
  • اگر شما یک سلّول زندهٔ اسفنج را در آب بیندازید، هفته آینده‌اش شما به اندازهٔ یک نخود اسفنج خواهید داشت! این امر قدرت رشد و ترمیم‌پذیری اسفنج را نشان می‌دهد.
  • اگر اسفنج را تکه‌تکه کنیم و حتی از صافی عبور دهیم، بعد از مدتی فقط یک اسفنج خواهیم دید؛ این نشان دهندهٔ قابلیت زیاد ترمیم اسفنج‌هاست.

طبقه‌بندی اسفنج‌ها[ویرایش]

اسفنج‌ها، از روی ظاهرشان به سه ردهٔ شیشه‌ای، آهکی و بزرگ تقسیم می‌شوند. اکثراً ردهٔ شاخی که به صورت شاخه شاخه است و اعضای آن با مرجان‌های شاخه‌ای اشتباه گرفته می‌شوند. رده جدید به نام اسفنج‌های همگون‌سوزنه نیز پذیرفته شده‌است.



ویکی‌پدیای انگلیسی.

  1. واژه‌های مصوّب فرهنگستان تا پایان دفتر یازدهم فرهنگ واژه‌های مصوّب
  2. شایان، زیست‌شناسی دورهٔ راهنمایی مرآت

دانشنامه رشد[پیوند مرده]

دانشنامه کودکان و نوجوانان آکسفورد. نشر نی. ۱۳۸۳. گروهی از مترجمان

Temporal range: Ediacaran–recent
A stove-pipe sponge
Scientific classification e
Kingdom: Animalia
Phylum: Porifera
Grant, 1836
Type species
Amphimedon queenslandica[1]

Parazoa/Ahistozoa (sans Placozoa)[2]

Sponges, the members of the phylum Porifera (/pəˈrɪfərə/; meaning "pore bearer"), are a basal Metazoa (animal) clade as a sister of the Diploblasts.[3][4][5][6][7] They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through them, consisting of jelly-like mesohyl sandwiched between two thin layers of cells. The branch of zoology that studies sponges is known as spongiology.[8]

Sponges have unspecialized cells that can transform into other types and that often migrate between the main cell layers and the mesohyl in the process. Sponges do not have nervous, digestive or circulatory systems. Instead, most rely on maintaining a constant water flow through their bodies to obtain food and oxygen and to remove wastes. Sponges were first to branch off the evolutionary tree from the common ancestor of all animals, making them the sister group of all other animals.[3]


The term sponge derives from the Ancient Greek word σπόγγος (spóngos).[9]


Sponge biodiversity and morphotypes at the lip of a wall site in 60 feet (20 m) of water. Included are the yellow tube sponge, Aplysina fistularis, the purple vase sponge, Niphates digitalis, the red encrusting sponge, Spirastrella coccinea [nl], and the gray rope sponge, Callyspongia sp.

Sponges are similar to other animals in that they are multicellular, heterotrophic, lack cell walls and produce sperm cells. Unlike other animals, they lack true tissues[10] and organs.[11] Some of them are radially symmetrical, but most are asymmetrical. The shapes of their bodies are adapted for maximal efficiency of water flow through the central cavity, where the water deposits nutrients and then leaves through a hole called the osculum. Many sponges have internal skeletons of spongin and/or spicules (skeletal-like fragments) of calcium carbonate or silicon dioxide.[10] All sponges are sessile aquatic animals, meaning that they attach to an underwater surface and remain fixed in place (i.e., do not travel). Although there are freshwater species, the great majority are marine (salt-water) species, ranging in habitat from tidal zones to depths exceeding 8,800 m (5.5 mi).

Although most of the approximately 5,000–10,000 known species of sponges feed on bacteria and other microscopic food in the water, some host photosynthesizing microorganisms as endosymbionts, and these alliances often produce more food and oxygen than they consume. A few species of sponges that live in food-poor environments have evolved as carnivores that prey mainly on small crustaceans.[12]

Most species use sexual reproduction, releasing sperm cells into the water to fertilize ova that in some species are released and in others are retained by the "mother." The fertilized eggs develop into larvae, which swim off in search of places to settle.[13] Sponges are known for regenerating from fragments that are broken off, although this only works if the fragments include the right types of cells. A few species reproduce by budding. When environmental conditions become less hospitable to the sponges, for example as temperatures drop, many freshwater species and a few marine ones produce gemmules, "survival pods" of unspecialized cells that remain dormant until conditions improve; they then either form completely new sponges or recolonize the skeletons of their parents.[14]

In most sponges, an internal gelatinous matrix called mesohyl functions as an endoskeleton, and it is the only skeleton in soft sponges that encrust such hard surfaces as rocks. More commonly, the mesohyl is stiffened by mineral spicules, by spongin fibers, or both. Demosponges use spongin; many species have silica spicules, whereas some species have calcium carbonate exoskeletons. Demosponges constitute about 90% of all known sponge species, including all freshwater ones, and they have the widest range of habitats. Calcareous sponges, which have calcium carbonate spicules and, in some species, calcium carbonate exoskeletons, are restricted to relatively shallow marine waters where production of calcium carbonate is easiest.[15] The fragile glass sponges, with "scaffolding" of silica spicules, are restricted to polar regions and the ocean depths where predators are rare. Fossils of all of these types have been found in rocks dated from 580 million years ago. In addition Archaeocyathids, whose fossils are common in rocks from 530 to 490 million years ago, are now regarded as a type of sponge.

The single-celled choanoflagellates resemble the choanocyte cells of sponges which are used to drive their water flow systems and capture most of their food. This along with phylogenetic studies of ribosomal molecules have been used as morphological evidence to suggest sponges are the sister group to the rest of animals.[16] Some studies have shown that sponges do not form a monophyletic group, in other words do not include all and only the descendants of a common ancestor. Recent phylogenetic analyses suggest that comb jellies rather than sponges are the sister group to the rest of animals.[17][18][19][20]


Distinguishing features

Sponges constitute the phylum Porifera, and have been defined as sessile metazoans (multicelled immobile animals) that have water intake and outlet openings connected by chambers lined with choanocytes, cells with whip-like flagella.[22] However, a few carnivorous sponges have lost these water flow systems and the choanocytes.[23][24] All known living sponges can remold their bodies, as most types of their cells can move within their bodies and a few can change from one type to another.[24][25]

Even if a few sponges are able to produce mucus – which acts as a microbial barrier in all other animals – no sponge with the ability to secrete a functional mucus layer has been recorded. Without such a mucus layer their living tissue is covered by a layer of microbial symbionts, which can contribute up to 40–50% of the sponge wet mass. This inability to prevent microbes from penetrating their porous tissue could be a major reason why they have never evolved a more complex anatomy.[26]

Like cnidarians (jellyfish, etc.) and ctenophores (comb jellies), and unlike all other known metazoans, sponges' bodies consist of a non-living jelly-like mass (mesoglea) sandwiched between two main layers of cells.[27][28] Cnidarians and ctenophores have simple nervous systems, and their cell layers are bound by internal connections and by being mounted on a basement membrane (thin fibrous mat, also known as "basal lamina").[28] Sponges have no nervous systems, their middle jelly-like layers have large and varied populations of cells, and some types of cells in their outer layers may move into the middle layer and change their functions.[25]

  Sponges[25][27] Cnidarians and ctenophores[28]
Nervous system No Yes, simple
Cells in each layer bound together No, except that Homoscleromorpha have basement membranes.[29] Yes: inter-cell connections; basement membranes
Number of cells in middle "jelly" layer Many Few
Cells in outer layers can move inwards and change functions Yes No

Basic structure

Cell types

    Water flow
Main cell types of Porifera[30]


Other types of cell live and move within the mesohyl:[25][27]

  • Collencytes are another type of collagen-producing cell.
  • Oocytes and spermatocytes are reproductive cells.

Many larval sponges possess neuron-less eyes that are based on cryptochromes. They mediate phototaxic behavior.[31]

Glass sponges' syncytia

    Water flow
    Main syncitium
    and collar bodies
    showing interior



Water flow and body structures

    Water flow
Porifera body structures[34]




Diagram of a syconoid sponge
Diagram of a syconoid sponge



    Archeocytes and other cells in
    Seabed / rock
    Water flow
Sponge with calcium carbonate skeleton[25]


In zoology a skeleton is any fairly rigid structure of an animal, irrespective of whether it has joints and irrespective of whether it is biomineralized. The mesohyl functions as an endoskeleton in most sponges, and is the only skeleton in soft sponges that encrust hard surfaces such as rocks. More commonly the mesohyl is stiffened by mineral spicules, by spongin fibers or both. Spicules, which are present in most but not all species,[37] may be made of silica or calcium carbonate, and vary in shape from simple rods to three-dimensional "stars" with up to six rays. Spicules are produced by sclerocyte cells,[25] and may be separate, connected by joints, or fused.[24]


Vital functions

Spongia officinalis, "the kitchen sponge", is dark grey when alive.



Respiration, feeding and excretion

Sponges do not have distinct circulatory, respiratory, digestive, and excretory systems – instead the water flow system supports all these functions. They filter food particles out of the water flowing through them. Particles larger than 50 micrometers cannot enter the ostia and pinacocytes consume them by phagocytosis (engulfing and internal digestion). Particles from 0.5 μm to 50 μm are trapped in the ostia, which taper from the outer to inner ends. These particles are consumed by pinacocytes or by archaeocytes which partially extrude themselves through the walls of the ostia. Bacteria-sized particles, below 0.5 micrometers, pass through the ostia and are caught and consumed by choanocytes.[25] Since the smallest particles are by far the most common, choanocytes typically capture 80% of a sponge's food supply.[38] Archaeocytes transport food packaged in vesicles from cells that directly digest food to those that do not. At least one species of sponge has internal fibers that function as tracks for use by nutrient-carrying archaeocytes,[25] and these tracks also move inert objects.[27]

Euplectella aspergillum, a glass sponge known as "Venus' flower basket"

It used to be claimed that glass sponges could live on nutrients dissolved in sea water and were very averse to silt.[39] However, a study in 2007 found no evidence of this and concluded that they extract bacteria and other micro-organisms from water very efficiently (about 79%) and process suspended sediment grains to extract such prey.[40] Collar bodies digest food and distribute it wrapped in vesicles that are transported by dynein "motor" molecules along bundles of microtubules that run throughout the syncytium.[25]


Carnivorous sponges

A few species that live in waters where the supply of food particles is very poor prey on crustaceans and other small animals. So far only 137 species have been discovered.[41] Most belong to the family Cladorhizidae, but a few members of the Guitarridae and Esperiopsidae are also carnivores.[42] In most cases little is known about how they actually capture prey, although some species are thought to use either sticky threads or hooked spicules.[42][43] Most carnivorous sponges live in deep waters, up to 8,840 m (5.49 mi),[44] and the development of deep-ocean exploration techniques is expected to lead to the discovery of several more.[25][42] However, one species has been found in Mediterranean caves at depths of 17–23 m (56–75 ft), alongside the more usual filter feeding sponges. The cave-dwelling predators capture crustaceans under 1 mm (0.039 in) long by entangling them with fine threads, digest them by enveloping them with further threads over the course of a few days, and then return to their normal shape; there is no evidence that they use venom.[44]

Most known carnivorous sponges have completely lost the water flow system and choanocytes. However, the genus Chondrocladia uses a highly modified water flow system to inflate balloon-like structures that are used for capturing prey.[42][45]


Freshwater sponges often host green algae as endosymbionts within archaeocytes and other cells, and benefit from nutrients produced by the algae. Many marine species host other photosynthesizing organisms, most commonly cyanobacteria but in some cases dinoflagellates. Symbiotic cyanobacteria may form a third of the total mass of living tissue in some sponges, and some sponges gain 48% to 80% of their energy supply from these micro-organisms.[25] In 2008 a University of Stuttgart team reported that spicules made of silica conduct light into the mesohyl, where the photosynthesizing endosymbionts live.[46] Sponges that host photosynthesizing organisms are most common in waters with relatively poor supplies of food particles, and often have leafy shapes that maximize the amount of sunlight they collect.[27]


"Immune" system




The freshwater sponge Spongilla lacustris

Sponges have three asexual methods of reproduction: after fragmentation; by budding; and by producing gemmules. Fragments of sponges may be detached by currents or waves. They use the mobility of their pinacocytes and choanocytes and reshaping of the mesohyl to re-attach themselves to a suitable surface and then rebuild themselves as small but functional sponges over the course of several days. The same capabilities enable sponges that have been squeezed through a fine cloth to regenerate.[47] A sponge fragment can only regenerate if it contains both collencytes to produce mesohyl and archeocytes to produce all the other cell types.[38] A very few species reproduce by budding.[48]

Gemmules are "survival pods" which a few marine sponges and many freshwater species produce by the thousands when dying and which some, mainly freshwater species, regularly produce in autumn. Spongocytes make gemmules by wrapping shells of spongin, often reinforced with spicules, round clusters of archeocytes that are full of nutrients.[49] Freshwater gemmules may also include phytosynthesizing symbionts.[50] The gemmules then become dormant, and in this state can survive cold, drying out, lack of oxygen and extreme variations in salinity.[25] Freshwater gemmules often do not revive until the temperature drops, stays cold for a few months and then reaches a near-"normal" level.[50] When a gemmule germinates, the archeocytes round the outside of the cluster transform into pinacocytes, a membrane over a pore in the shell bursts, the cluster of cells slowly emerges, and most of the remaining archeocytes transform into other cell types needed to make a functioning sponge. Gemmules from the same species but different individuals can join forces to form one sponge.[51] Some gemmules are retained within the parent sponge, and in spring it can be difficult to tell whether an old sponge has revived or been "recolonized" by its own gemmules.[50]





Life cycle


Coordination of activities

Adult sponges lack neurons or any other kind of nervous tissue. However, most species have the ability to perform movements that are coordinated all over their bodies, mainly contractions of the pinacocytes, squeezing the water channels and thus expelling excess sediment and other substances that may cause blockages. Some species can contract the osculum independently of the rest of the body. Sponges may also contract in order to reduce the area that is vulnerable to attack by predators. In cases where two sponges are fused, for example if there is a large but still unseparated bud, these contraction waves slowly become coordinated in both of the "Siamese twins". The coordinating mechanism is unknown, but may involve chemicals similar to neurotransmitters.[54] However, glass sponges rapidly transmit electrical impulses through all parts of the syncytium, and use this to halt the motion of their flagella if the incoming water contains toxins or excessive sediment.[25] Myocytes are thought to be responsible for closing the osculum and for transmitting signals between different parts of the body.[27]





Sponges are worldwide in their distribution, living in a wide range of ocean habitats, from the polar regions to the tropics.[38] Most live in quiet, clear waters, because sediment stirred up by waves or currents would block their pores, making it difficult for them to feed and breathe.[39] The greatest numbers of sponges are usually found on firm surfaces such as rocks, but some sponges can attach themselves to soft sediment by means of a root-like base.[56]


The different classes of sponge live in different ranges of habitat:

  Water type[27] Depth[27] Type of surface[27]
Calcarea Marine less than 100 m (330 ft) Hard
Glass sponges Marine Deep Soft or firm sediment
Demosponges Marine, brackish; and about 150 freshwater species[25] Inter-tidal to abyssal;[27] a carnivorous demosponge has been found at 8,840 m (5.49 mi)[44] Any

As primary producers




Many sponges shed spicules, forming a dense carpet several meters deep that keeps away echinoderms which would otherwise prey on the sponges.[38] They also produce toxins that prevent other sessile organisms such as bryozoans or sea squirts from growing on or near them, making sponges very effective competitors for living space. One of many examples includes ageliferin.

A few species, the Caribbean fire sponge Tedania ignis, cause a severe rash in humans who handle them.[25] Turtles and some fish feed mainly on sponges. It is often said that sponges produce chemical defenses against such predators.[25] However, experiments have been unable to establish a relationship between the toxicity of chemicals produced by sponges and how they taste to fish, which would diminish the usefulness of chemical defenses as deterrents. Predation by fish may even help to spread sponges by detaching fragments.[27] However, some studies have shown fish showing a preference for non chemically defended sponges,[59] and another study found that high levels of coral predation did predict the presence of chemically defended species.[60]

Glass sponges produce no toxic chemicals, and live in very deep water where predators are rare.[39]




The Caribbean chicken-liver sponge Chondrilla nucula secretes toxins that kill coral polyps, allowing the sponges to grow over the coral skeletons.[25] Others, especially in the family Clionaidae, use corrosive substances secreted by their archeocytes to tunnel into rocks, corals and the shells of dead mollusks.[25] Sponges may remove up to 1 m (3.3 ft) per year from reefs, creating visible notches just below low-tide level.[38]


Caribbean sponges of the genus Aplysina suffer from Aplysina red band syndrome. This causes Aplysina to develop one or more rust-colored bands, sometimes with adjacent bands of necrotic tissue. These lesions may completely encircle branches of the sponge. The disease appears to be contagious and impacts approximately 10 percent of A. cauliformis on Bahamian reefs.[62] The rust-colored bands are caused by a cyanobacterium, but it is unknown whether this organism actually causes the disease.[62][63]

Collaboration with other organisms



Systematics and evolutionary history


Linnaeus, who classified most kinds of sessile animals as belonging to the order Zoophyta in the class Vermes, mistakenly identified the genus Spongia as plants in the order Algae.[67] For a long time thereafter sponges were assigned to a separate subkingdom, Parazoa ("beside the animals"), separate from the Eumetazoa which formed the rest of the kingdom Animalia.[68] They have been regarded as a paraphyletic phylum, from which the higher animals have evolved.[69] Other research indicates Porifera is monophyletic.[70]

The phylum Porifera is further divided into classes mainly according to the composition of their skeletons:[24][38]

  • Hexactinellida (glass sponges) have silicate spicules, the largest of which have six rays and may be individual or fused.[24] The main components of their bodies are syncytia in which large numbers of cell share a single external membrane.[38]
  • Most Demospongiae have silicate spicules or spongin fibers or both within their soft tissues. However, a few also have massive external skeletons made of aragonite, another form of calcium carbonate.[24][38] All the cells have a single nucleus and membrane.[38]
  • Archeocyatha are known only as fossils from the Cambrian period.[68]

In the 1970s, sponges with massive calcium carbonate skeletons were assigned to a separate class, Sclerospongiae, otherwise known as "coralline sponges".[71] However, in the 1980s it was found that these were all members of either the Calcarea or the Demospongiae.[72]

So far scientific publications have identified about 9,000 poriferan species,[38] of which: about 400 are glass sponges; about 500 are calcareous species; and the rest are demosponges.[25] However, some types of habitat, vertical rock and cave walls and galleries in rock and coral boulders, have been investigated very little, even in shallow seas.[38]


Sponges were traditionally distributed in three classes: calcareous sponges (Calcarea), glass sponges (Hexactinellida) and demosponges (Demospongiae). However, studies have shown that the Homoscleromorpha, a group thought to belong to the Demospongiae, is actually phylogenetically well separated.[73] Therefore, they have recently been recognized as the fourth class of sponges.[74][75]

Sponges are divided into classes mainly according to the composition of their skeletons:[27]

  Type of cells[27] Spicules[27] Spongin fibers[27] Massive exoskeleton[38] Body form[27]
Calcarea Single nucleus, single external membrane Calcite
May be individual or large masses
Never Common.
Made of calcite if present.
Asconoid, syconoid, leuconoid or solenoid[76]
Hexactinellida Mostly syncytia in all species Silica
May be individual or fused
Never Never Leuconoid
Made of aragonite if present.[24][38]

Fossil record

Raphidonema faringdonense, a fossil sponge from the Cretaceous of England
Archaeocyathid structure

Although molecular clocks and biomarkers suggest sponges existed well before the Cambrian explosion of life, silica spicules like those of demosponges are absent from the fossil record until the Cambrian.[77] One unsubstantiated report exists of spicules in rocks dated around 750 million years ago.[78] Well-preserved fossil sponges from about 580 million years ago in the Ediacaran period have been found in the Doushantuo Formation. These fossils, which include spicules, pinacocytes, porocytes, archeocytes, sclerocytes and the internal cavity, have been classified as demosponges. Fossils of glass sponges have been found from around 540 million years ago in rocks in Australia, China and Mongolia.[79] Early Cambrian sponges from Mexico belonging to the genus Kiwetinokia show evidence of fusion of several smaller spicules to form a single large spicule.[80] Calcium carbonate spicules of calcareous sponges have been found in Early Cambrian rocks from about 530 to 523 million years ago in Australia. Other probable demosponges have been found in the Early Cambrian Chengjiang fauna, from 525 to 520 million years ago.[81] Freshwater sponges appear to be much younger, as the earliest known fossils date from the Mid-Eocene period about 48 to 40 million years ago.[79] Although about 90% of modern sponges are demosponges, fossilized remains of this type are less common than those of other types because their skeletons are composed of relatively soft spongin that does not fossilize well.[82] Earliest sponge symbionts are known from the early Silurian.[83]

A chemical tracer is 24-isopropylcholestane, which is a stable derivative of 24-isopropylcholesterol, which is said to be produced by demosponges but not by eumetazoans ("true animals", i.e. cnidarians and bilaterians). Since choanoflagellates are thought to be animals' closest single-celled relatives, a team of scientists examined the biochemistry and genes of one choanoflagellate species. They concluded that this species could not produce 24-isopropylcholesterol but that investigation of a wider range of choanoflagellates would be necessary in order to prove that the fossil 24-isopropylcholestane could only have been produced by demosponges.[84] Although a previous publication reported traces of the chemical 24-isopropylcholestane in ancient rocks dating to 1,800 million years ago,[85] recent research using a much more accurately dated rock series has revealed that these biomarkers only appear before the end of the Marinoan glaciation approximately 635 million years ago,[86] and that "Biomarker analysis has yet to reveal any convincing evidence for ancient sponges pre-dating the first globally extensive Neoproterozoic glacial episode (the Sturtian, ~713 million years ago in Oman)". While it has been argued that this 'sponge biomarker' could have originated from marine algae, recent research suggests that the algae's ability to produce this biomarker evolved only in the Carboniferous; as such, the biomarker remains strongly supportive of the presence of demosponges in the Cryogenian.[87][88][89]


= skin
= flesh
Halkieriid sclerite structure[90]


Relationships to other animal groups

A choanoflagellate




Glass sponges


Calcareous sponges


Comb jellies


(jellyfish, etc.)

Simplified family tree showing calcareous sponges
as closest to more complex animals[93]




Most demosponges

Calcareous sponges



(jellyfish, etc.)

Other metazoans

Simplified family tree showing Homoscleromorpha
as closest to more complex animals[94]






Notable spongiologists


Sponges made of sponge gourd for sale alongside sponges of animal origin (Spice Bazaar at Istanbul, Turkey).

By dolphins


By humans

Natural sponges in Tarpon Springs, Florida
Display of natural sponges for sale on Kalymnos in Greece


The calcium carbonate or silica spicules of most sponge genera make them too rough for most uses, but two genera, Hippospongia and Spongia, have soft, entirely fibrous skeletons.[105] Early Europeans used soft sponges for many purposes, including padding for helmets, portable drinking utensils and municipal water filters. Until the invention of synthetic sponges, they were used as cleaning tools, applicators for paints and ceramic glazes and discreet contraceptives. However, by the mid-20th century, over-fishing brought both the animals and the industry close to extinction.[106] See also sponge diving.

Many objects with sponge-like textures are now made of substances not derived from poriferans. Synthetic sponges include personal and household cleaning tools, breast implants,[107] and contraceptive sponges.[108] Typical materials used are cellulose foam, polyurethane foam, and less frequently, silicone foam.


Antibiotic compounds

Sponges have medicinal potential due to the presence in sponges themselves or their microbial symbionts of chemicals that may be used to control viruses, bacteria, tumors and fungi.[110][111]

Other biologically active compounds

Halichondria produces the eribulin precursor halichondrin B


See also


  1. ^ Srivastava M, Simakov O, Chapman J, Fahey B, Gauthier ME, Mitros T, et al. (August 2010). "The Amphimedon queenslandica genome and the evolution of animal complexity". Nature. 466 (7307): 720–6. Bibcode:2010Natur.466..720S. doi:10.1038/nature09201. PMC 3130542. PMID 20686567.
  2. ^ Pajdzińska A (2018). "Animals die more shallowly: they aren't deceased, they're dead. Animals in the polish linguistic worldview and in contemporary life sciences" (PDF). Ethnolinguistic. 29: 147–161. doi:10.17951/et.2017.29.135.
  3. ^ a b Feuda R, Dohrmann M, Pett W, Philippe H, Rota-Stabelli O, Lartillot N, et al. (December 2017). "Improved Modeling of Compositional Heterogeneity Supports Sponges as Sister to All Other Animals". Current Biology. 27 (24): 3864–3870.e4. doi:10.1016/j.cub.2017.11.008. PMID 29199080.
  4. ^ Pisani D, Pett W, Dohrmann M, Feuda R, Rota-Stabelli O, Philippe H, et al. (December 2015). "Genomic data do not support comb jellies as the sister group to all other animals". Proceedings of the National Academy of Sciences of the United States of America. 112 (50): 15402–7. Bibcode:2015PNAS..11215402P. doi:10.1073/pnas.1518127112. PMC 4687580. PMID 26621703.
  5. ^ a b Simion P, Philippe H, Baurain D, Jager M, Richter DJ, Di Franco A, et al. (April 2017). "A Large and Consistent Phylogenomic Dataset Supports Sponges as the Sister Group to All Other Animals" (PDF). Current Biology. 27 (7): 958–967. Bibcode:1996CBio....6.1213A. doi:10.1016/j.cub.2017.02.031. PMID 28318975.
  7. ^ Laumer CE, Gruber-Vodicka H, Hadfield MG, Pearse VB, Riesgo A, Marioni JC, Giribet G (2017-10-11). "Placozoans are eumetazoans related to Cnidaria". bioRxiv 200972.
  8. ^ "Spongiology". Merriam-Webster Dictionary. Retrieved 27 December 2017.
  10. ^ a b Hooper, John (2018). "Structure of Sponges". Queensland Museum. Retrieved 27 September 2019.
  11. ^ Thacker, Robert W; Diaz, Maria Christina (8 September 2014). "The Porifera Ontology (PORO): enhancing sponge systematics with an anatomy ontology". J Biomed Semantics. 5 (39): 39. doi:10.1186/2041-1480-5-39. PMC 4177528. PMID 25276334.
  12. ^ Vacelet & Duport 2004, pp. 179–190.
  13. ^ Bergquist 1978, pp. 183–185.
  14. ^ Bergquist 1978, pp. 120–127.
  15. ^ Bergquist 1978, p. 179.
  16. ^ Collins AG (December 1998). "Evaluating multiple alternative hypotheses for the origin of Bilateria: an analysis of 18S rRNA molecular evidence". Proceedings of the National Academy of Sciences of the United States of America. 95 (26): 15458–63. Bibcode:1998PNAS...9515458C. doi:10.1073/pnas.95.26.15458. PMC 28064. PMID 9860990.
  17. ^ a b Dunn CW, Hejnol A, Matus DQ, Pang K, Browne WE, Smith SA, et al. (April 2008). "Broad phylogenomic sampling improves resolution of the animal tree of life". Nature. 452 (7188): 745–9. Bibcode:2008Natur.452..745D. doi:10.1038/nature06614. PMID 18322464.
  18. ^ Hejnol A, Obst M, Stamatakis A, Ott M, Rouse GW, Edgecombe GD, et al. (December 2009). "Assessing the root of bilaterian animals with scalable phylogenomic methods". Proceedings. Biological Sciences. 276 (1677): 4261–70. doi:10.1098/rspb.2009.0896. PMC 2817096. PMID 19759036.
  19. ^ Ryan JF, Pang K, Schnitzler CE, Nguyen AD, Moreland RT, Simmons DK, et al. (December 2013). "The genome of the ctenophore Mnemiopsis leidyi and its implications for cell type evolution". Science. 342 (6164): 1242592. doi:10.1126/science.1242592. PMC 3920664. PMID 24337300.
  20. ^ Moroz LL, Kocot KM, Citarella MR, Dosung S, Norekian TP, Povolotskaya IS, et al. (June 2014). "The ctenophore genome and the evolutionary origins of neural systems". Nature. 510 (7503): 109–14. Bibcode:2014Natur.510..109M. doi:10.1038/nature13400. PMC 4337882. PMID 24847885.
  21. ^ a b Krützen M, Mann J, Heithaus MR, Connor RC, Bejder L, Sherwin WB (June 2005). "Cultural transmission of tool use in bottlenose dolphins". Proceedings of the National Academy of Sciences of the United States of America. 102 (25): 8939–43. Bibcode:2005PNAS..102.8939K. doi:10.1073/pnas.0500232102. PMC 1157020. PMID 15947077.
  22. ^ Bergquist 1978, p. 29.
  23. ^ Bergquist 1978, p. 39.
  24. ^ a b c d e f g Hooper JN, Van Soest RW, Debrenne F (2002). "Phylum Porifera Grant, 1836". In Hooper JN, Van Soest RW (eds.). Systema Porifera: A Guide to the Classification of Sponges. New York: Kluwer Academic/Plenum. pp. 9–14. ISBN 978-0-306-47260-2.
  26. ^ Bakshani CR, Morales-Garcia AL, Althaus M, Wilcox MD, Pearson JP, Bythell JC, Burgess JG (2018-07-04). "Evolutionary conservation of the antimicrobial function of mucus: a first defence against infection". NPJ Biofilms and Microbiomes. 4 (1): 14. doi:10.1038/s41522-018-0057-2. PMC 6031612. PMID 30002868.
  27. ^ a b c d e f g h i j k l m n o p q r s t Bergquist PR (1998). "Porifera". In Anderson DT (ed.). Invertebrate Zoology. Oxford University Press. pp. 10–27. ISBN 978-0-19-551368-4.
  28. ^ a b c Hinde RT (1998). "The Cnidaria and Ctenophora". In Anderson DT (ed.). Invertebrate Zoology. Oxford University Press. pp. 28–57. ISBN 978-0-19-551368-4.
  29. ^ a b Exposito JY, Cluzel C, Garrone R, Lethias C (November 2002). "Evolution of collagens". The Anatomical Record. 268 (3): 302–16. doi:10.1002/ar.10162. PMID 12382326.
  30. ^ Ruppert EE, Fox RS, Barnes RD (2004). Invertebrate Zoology (7th ed.). Brooks / Cole. p. 82. ISBN 978-0-03-025982-1.
  31. ^ Rivera AS, Ozturk N, Fahey B, Plachetzki DC, Degnan BM, Sancar A, Oakley TH (April 2012). "Blue-light-receptive cryptochrome is expressed in a sponge eye lacking neurons and opsin". The Journal of Experimental Biology. 215 (Pt 8): 1278–86. doi:10.1242/jeb.067140. PMC 3309880. PMID 22442365.
  32. ^ Ruppert EE, Fox RS, Barnes RD (2004). Invertebrate Zoology (7th ed.). Brooks / Cole. p. 83. ISBN 978-0-03-025982-1. Fig. 5-7
  33. ^ a b Leys SP (February 2003). "The significance of syncytial tissues for the position of the hexactinellida in the metazoa". Integrative and Comparative Biology. 43 (1): 19–27. doi:10.1093/icb/43.1.19. PMID 21680406.
  34. ^ Ruppert EE, Fox RS, Barnes RD (2004). Invertebrate Zoology (7th ed.). Brooks / Cole. p. 78. ISBN 978-0-03-025982-1.
  35. ^ Ruppert, Fox & Barnes 2004, p. 83.
  36. ^ Hickman CP, Roberts LS, Larson A (2001). Integrated Principles of Zoology (11th ed.). New York: McGraw-Hill. p. 247. ISBN 978-0-07-290961-6.
  37. ^ Halisarca dujardini - Marine Species Identification Portal
  38. ^ a b c d e f g h i j k l m n o Bergquist PR (2001). "Porifera (Sponges)". Encyclopedia of Life Sciences. John Wiley & Sons, Ltd. doi:10.1038/npg.els.0001582. ISBN 978-0470016176.
  40. ^ Yahel G, Whitney F, Reiswig HM, Eerkes-Medrano DI, Leys SP (2007). "In situ feeding and metabolism of glass sponges (Hexactinellida, Porifera) studied in a deep temperate fjord with a remotely operated submersible". Limnology and Oceanography. 52 (1): 428–440. Bibcode:2007LimOc..52..428Y. CiteSeerX doi:10.4319/lo.2007.52.1.0428.
  42. ^ a b c d Vacelet J (2008). "A new genus of carnivorous sponges (Porifera: Poecilosclerida, Cladorhizidae) from the deep N-E Pacific, and remarks on the genus Neocladia" (PDF). Zootaxa. 1752: 57–65. doi:10.11646/zootaxa.1752.1.3. Retrieved 2008-10-31.
  44. ^ a b c Vacelet J, Boury-Esnault N (1995). "Carnivorous sponges". Nature. 373 (6512): 333–335. Bibcode:1995Natur.373..333V. doi:10.1038/373333a0.
  45. ^ Vacelet J, Kelly M (2008). "New species from the deep Pacific suggest that carnivorous sponges date back to the Early Jurassic". Nature Precedings. doi:10.1038/npre.2008.2327.1.
  46. ^ Brümmer F, Pfannkuchen M, Baltz A, Hauser T, Thiel V (2008). "Light inside sponges". Journal of Experimental Marine Biology and Ecology. 367 (2): 61–64. doi:10.1016/j.jembe.2008.06.036. Lay summaryBBC News.
  47. ^ Ruppert, Fox & Barnes 2004, p. 239.
  48. ^ Ruppert, Fox & Barnes 2004, pp. 90–94.
  49. ^ Ruppert, Fox & Barnes 2004, pp. 87–88.
  50. ^ a b c Smith DG, Pennak RW (2001). Pennak's Freshwater Invertebrates of the United States: Porifera to Crustacea (4 ed.). John Wiley and Sons. pp. 47–50. ISBN 978-0-471-35837-4.
  51. ^ Ruppert, Fox & Barnes 2004, pp. 89–90.
  52. ^ a b Ruppert, Fox & Barnes 2004, p. 77.
  53. ^ Leys SP, Cheung E, Boury-Esnault N (April 2006). "Embryogenesis in the glass sponge Oopsacas minuta: Formation of syncytia by fusion of blastomeres". Integrative and Comparative Biology. 46 (2): 104–17. doi:10.1093/icb/icj016. PMID 21672727.
  54. ^ Nickel M (December 2004). "Kinetics and rhythm of body contractions in the sponge Tethya wilhelma (Porifera: Demospongiae)". The Journal of Experimental Biology. 207 (Pt 26): 4515–24. doi:10.1242/jeb.01289. PMID 15579547.
  55. ^ Sakarya O, Armstrong KA, Adamska M, Adamski M, Wang IF, Tidor B, et al. (June 2007). "A post-synaptic scaffold at the origin of the animal kingdom". PLOS ONE. 2 (6): e506. Bibcode:2007PLoSO...2..506S. doi:10.1371/journal.pone.0000506. PMC 1876816. PMID 17551586.
  56. ^ Weaver JC, Aizenberg J, Fantner GE, Kisailus D, Woesz A, Allen P, et al. (April 2007). "Hierarchical assembly of the siliceous skeletal lattice of the hexactinellid sponge Euplectella aspergillum". Journal of Structural Biology. 158 (1): 93–106. doi:10.1016/j.jsb.2006.10.027. PMID 17175169.
  57. ^ Ruzicka R, Gleason DF (January 2008). "Latitudinal variation in spongivorous fishes and the effectiveness of sponge chemical defenses" (PDF). Oecologia. 154 (4): 785–94. Bibcode:2008Oecol.154..785R. doi:10.1007/s00442-007-0874-0. PMID 17960425. Archived from the original (PDF) on 2008-10-06.
  58. ^ Gage & Tyler 1996, pp. 91–93
  59. ^ Dunlap M, Pawlik JR (1996). "Video-monitored predation by Caribbean reef fishes on an array of mangrove and reef sponges". Marine Biology. 126 (1): 117–123. doi:10.1007/bf00571383. ISSN 0025-3162.
  60. ^ Loh TL, Pawlik JR (March 2014). "Chemical defenses and resource trade-offs structure sponge communities on Caribbean coral reefs". Proceedings of the National Academy of Sciences of the United States of America. 111 (11): 4151–6. Bibcode:2014PNAS..111.4151L. doi:10.1073/pnas.1321626111. PMC 3964098. PMID 24567392.
  61. ^ Piper 2007, p. 148.
  62. ^ a b Gochfeld DJ, Easson CG, Slattery M, Thacker RW, Olson JB (2012). Steller D, Lobel L (eds.). "Population Dynamics of a Sponge Disease on Caribbean Reefs". Diving for Science 2012. Proceedings of the American Academy of Underwater Sciences 31st Symposium. Retrieved 2013-11-17.
  63. ^ Olson JB, Gochfeld DJ, Slattery M (July 2006). "Aplysina red band syndrome: a new threat to Caribbean sponges" (PDF). Diseases of Aquatic Organisms. 71 (2): 163–8. doi:10.3354/dao071163. PMID 16956064. Lay summaryPractical Fishkeeping.
  64. ^ Wulff JL (June 2008). "Collaboration among sponge species increases sponge diversity and abundance in a seagrass meadow". Marine Ecology. 29 (2): 193–204. Bibcode:2008MarEc..29..193W. doi:10.1111/j.1439-0485.2008.00224.x.
  65. ^ Duffy JE (1996). "Species boundaries, specialization, and the radiation of sponge-dwelling alpheid shrimp" (PDF). Biological Journal of the Linnean Society. 58 (3): 307–324. doi:10.1111/j.1095-8312.1996.tb01437.x. Archived from the original (PDF) on August 3, 2010.
  66. ^ Murphy 2002, p. 51.
  69. ^ Sperling EA, Pisani D, Peterson KJ (January 1, 2007). "Poriferan paraphyly and its implications for Precambrian palaeobiology" (PDF). Geological Society, London, Special Publications. 286 (1): 355–368. Bibcode:2007GSLSP.286..355S. doi:10.1144/SP286.25. Archived from the original (PDF) on May 9, 2009. Retrieved 2012-08-22.
  70. ^ Whelan NV, Kocot KM, Moroz LL, Halanych KM (May 2015). "Error, signal, and the placement of Ctenophora sister to all other animals". Proceedings of the National Academy of Sciences of the United States of America. 112 (18): 5773–8. Bibcode:2015PNAS..112.5773W. doi:10.1073/pnas.1503453112. PMC 4426464. PMID 25902535.
  71. ^ Hartman WD, Goreau TF (1970). "Jamaican coralline sponges: Their morphology, ecology and fossil relatives". Symposium of the Zoological Society of London. 25: 205–243. (cited by MGG.rsmas.miami.edu).
  72. ^ Vacelet J (1985). "Coralline sponges and the evolution of the Porifera". In Conway Morris S, George JD, Gibson R, Platt HM (eds.). The Origins and Relationships of Lower Invertebrates. Oxford University Press. pp. 1–13. ISBN 978-0-19-857181-0.
  73. ^ Bergquist 1978, pp. 153–154.
  74. ^ Gazave E, Lapébie P, Renard E, Vacelet J, Rocher C, Ereskovsky AV, Lavrov DV, Borchiellini C (December 2010). "Molecular phylogeny restores the supra-generic subdivision of homoscleromorph sponges (Porifera, Homoscleromorpha)". PLOS ONE. 5 (12): e14290. Bibcode:2010PLoSO...514290G. doi:10.1371/journal.pone.0014290. PMC 3001884. PMID 21179486.
  75. ^ Gazave E, Lapébie P, Ereskovsky AV, Vacelet J, Renard E, Cárdenas P, Borchiellini C (May 2012). "No longer Demospongiae: Homoscleromorpha formal nomination as a fourth class of Porifera" (PDF). Hydrobiologia. 687: 3–10. doi:10.1007/s10750-011-0842-x.
  76. ^ Cavalcanti FF, Klautau M (2011). "Solenoid: a new aquiferous system to Porifera". Zoomorphology. 130 (4): 255–260. doi:10.1007/s00435-011-0139-7.
  77. ^ Sperling EA, Robinson JM, Pisani D, Peterson KJ (January 2010). "Where's the glass? Biomarkers, molecular clocks, and microRNAs suggest a 200-Myr missing Precambrian fossil record of siliceous sponge spicules". Geobiology. 8 (1): 24–36. doi:10.1111/j.1472-4669.2009.00225.x. PMID 19929965.
  78. ^ Reitner J, Wörheide G (2002). "Non-Lithistid Fossil Demospongiae – Origins of their Palaeobiodiversity and Highlights in History of Preservation". In Hooper JN, Van Soest RW (eds.). Systema Porifera: A Guide to the Classification of Sponges (PDF). New York: Kluwer Academic Plenum. Retrieved November 4, 2008.
  79. ^ a b c Müller WE, Li J, Schröder HC, Qiao L, Wang X (2007). "The unique skeleton of siliceous sponges (Porifera; Hexactinellida and Demospongiae) that evolved first from the Urmetazoa during the Proterozoic: a review". Biogeosciences. 4 (2): 219–232. doi:10.5194/bg-4-219-2007.
  81. ^ a b Li CW, Chen JY, Hua TE (February 1998). "Precambrian sponges with cellular structures". Science. 279 (5352): 879–82. Bibcode:1998Sci...279..879L. doi:10.1126/science.279.5352.879. PMID 9452391.
  83. ^ Vinn O, Wilson MA, Toom U, Mõtus MA (2015). "Earliest known rugosan-stromatoporoid symbiosis from the Llandovery of Estonia (Baltica)". Palaeogeography, Palaeoclimatology, Palaeoecology. 31: 1–5. Bibcode:2015PPP...431....1V. doi:10.1016/j.palaeo.2015.04.023. Retrieved 2015-06-18.
  84. ^ Kodner RB, Summons RE, Pearson A, King N, Knoll AH (July 2008). "Sterols in a unicellular relative of the metazoans". Proceedings of the National Academy of Sciences of the United States of America. 105 (29): 9897–902. Bibcode:2008PNAS..105.9897K. doi:10.1073/pnas.0803975105. PMC 2481317. PMID 18632573.
  85. ^ Nichols S, Wörheide G (April 2005). "Sponges: new views of old animals". Integrative and Comparative Biology. 45 (2): 333–4. CiteSeerX doi:10.1093/icb/45.2.333. PMID 21676777.
  86. ^ Love GD, Grosjean E, Stalvies C, Fike DA, Grotzinger JP, Bradley AS, Kelly AE, Bhatia M, Meredith W, Snape CE, Bowring SA, Condon DJ, Summons RE (February 2009). "Fossil steroids record the appearance of Demospongiae during the Cryogenian period" (PDF). Nature. 457 (7230): 718–21. Bibcode:2009Natur.457..718L. doi:10.1038/nature07673. PMID 19194449.
  87. ^ Antcliffe JB (2013). Stouge S (ed.). "Questioning the evidence of organic compounds called sponge biomarkers". Palaeontology. 56: 917–925. doi:10.1111/pala.12030.
  88. ^ Gold DA (Jun 29, 2018). "The slow rise of complex life as revealed through biomarker genetics". Emerging Topics in Life Sciences. 2 (2): 191–199. doi:10.1042/ETLS20170150.
  89. ^ Gold DA, Grabenstatter J, de Mendoza A, Riesgo A, Ruiz-Trillo I, Summons RE (March 2016). "Sterol and genomic analyses validate the sponge biomarker hypothesis". Proceedings of the National Academy of Sciences of the United States of America. 113 (10): 2684–9. Bibcode:2016PNAS..113.2684G. doi:10.1073/pnas.1512614113. PMC 4790988. PMID 26903629.
  90. ^ a b Porter SM (2008). "Skeletal microstructure indicates Chancelloriids and Halkieriids are closely related". Palaeontology. 51 (4): 865–879. doi:10.1111/j.1475-4983.2008.00792.x.
  91. ^ Butterfield NJ, Nicholas CJ (1996). "Burgess Shale-type preservation of both non-mineralizing and "shelly" Cambrian organisms from the Mackenzie Mountains, northwestern Canada". Journal of Paleontology. 70 (6): 893–899. doi:10.1017/S0022336000038579. JSTOR 1306492.
  92. ^ Janussen D, Steiner M, Zhu MY (2002). "New Well-preserved Scleritomes of Chancelloridae from the Early Cambrian Yuanshan Formation (Chengjiang, China) and the Middle Cambrian Wheeler Shale (Utah, USA) and paleobiological implications". Journal of Paleontology. 76 (4): 596–606. doi:10.1666/0022-3360(2002)076<0596:NWPSOC>2.0.CO;2. Free full text without images at Janussen D (2002). "(as above)". Journal of Paleontology. Archived from the original on December 10, 2008. Retrieved 2008-08-04.
  93. ^ a b c Borchiellini C, Manuel M, Alivon E, Boury-Esnault N, Vacelet J, Le Parco Y (January 2001). "Sponge paraphyly and the origin of Metazoa". Journal of Evolutionary Biology. 14 (1): 171–179. doi:10.1046/j.1420-9101.2001.00244.x. PMID 29280585.
  94. ^ a b Sperling EA, Pisani D, Peterson KJ (2007). "Poriferan paraphyly and its implications for Precambrian paleobiology" (PDF). Journal of the Geological Society of London. 286: 355–368. Bibcode:2007GSLSP.286..355S. doi:10.1144/SP286.25. Archived from the original (PDF) on May 9, 2009. Retrieved 2008-11-04.
  95. ^ Medina M, Collins AG, Silberman JD, Sogin ML (August 2001). "Evaluating hypotheses of basal animal phylogeny using complete sequences of large and small subunit rRNA". Proceedings of the National Academy of Sciences of the United States of America. 98 (17): 9707–12. Bibcode:2001PNAS...98.9707M. doi:10.1073/pnas.171316998. PMC 55517. PMID 11504944.
  96. ^ Schierwater B, Eitel M, Jakob W, Osigus HJ, Hadrys H, Dellaporta SL, et al. (January 2009). "Concatenated analysis sheds light on early metazoan evolution and fuels a modern "urmetazoon" hypothesis". PLoS Biology. 7 (1): e20. doi:10.1371/journal.pbio.1000020. PMC 2631068. PMID 19175291.
  97. ^ a b c d Schoenberg CH (17 June 2015). Van Soest RW, et al. (eds.). "Who is who in sponge science". The World Porifera Database. doi:10.13140/RG.2.1.1499.1526.
  98. ^ "Chemistry and the Natural Defences of Coral – ITW with Bernard Banaigs". Tara Expeditions Foundation. Retrieved 15 April 2018.
  99. ^ "Willard D. Hartman". Invertebrate Zoology : Collections : Yale Peabody Museum of Natural History. 7 December 2010. Retrieved 15 April 2018.
  100. ^ Hooper JN, van Soest RW (2012). Systema Porifera: A Guide to the Classification of Sponges. Springer Science & Business Media. p. 30. ISBN 978-1-4615-0747-5.
  101. ^ Annals & Magazine of Natural History. Taylor & Francis, Limited. 1875. p. 377.
  102. ^ Pansini M (2004). Sponge Science in the New Millennium: Papers Contributed to the VI International Sponge Conference, Rapallo (Italy), 29 September-5 October 2002. Universitá di Genova. p. 89.
  103. ^ "THE VOYAGE OF H.M.S. CHALLENGER". www.19thcenturyscience.org. Zoology, Part LIX, Volume XX. 1887.
  104. ^ Smolker RA, Richards AF, Connor RC, Mann J, Berggren P (1997). "Sponge-carrying by Indian Ocean bottlenose dolphins: Possible tool-use by a delphinid". Ethology. 103 (6): 454–465. doi:10.1111/j.1439-0310.1997.tb00160.x. hdl:2027.42/71936.
  105. ^ Bergquist 1978, p. 88.
  106. ^ McClenachan L (2008). "Social conflict, Over-fishing and Disease in the Florida Sponge Fishery, 1849–1939". In Starkey DJ, Holm P, Barnard M (eds.). Oceans Past: Management Insights from the History of Marine Animal Populations. Earthscan. pp. 25–27. ISBN 978-1-84407-527-0.
  107. ^ Jacobson N (2000). Cleavage. Rutgers University Press. p. 62. ISBN 978-0-8135-2715-4.
  109. ^ Porterfield WM (1955). "Loofah — The sponge gourd". Economic Botany. 9 (3): 211–223. doi:10.1007/BF02859814.
  110. ^ Imhoff JF, Stöhr R (2003). "Sponge-Associated Bacteria". In Müller WE (ed.). Sponges (Porifera): Porifera. Springer. pp. 43–44. ISBN 978-3-540-00968-9.
  111. ^ Teeyapant R, Woerdenbag HJ, Kreis P, Hacker J, Wray V, Witte L, Proksch P (1993). "Antibiotic and cytotoxic activity of brominated compounds from the marine sponge Verongia aerophoba". Zeitschrift für Naturforschung C. 48 (11–12): 939–45. doi:10.1515/znc-1993-11-1218. PMID 8297426.
  112. ^ Takeuchi S, Ishibashi M, Kobayashi J, Plakoridine A (1994). "Plakoridine A, a new tyramine-containing pyrrolidine alkaloid from the Okinawan marine sponge Plakortis sp". Journal of Organic Chemistry. 59 (13): 3712–3713. doi:10.1021/jo00092a039.
  113. ^ Etchells LL, Sardarian A, Whitehead RC (18 April 2005). "A synthetic approach to the plakoridines modeled on a biogenetic theory". Tetrahedron Letters. 46 (16): 2803–2807. doi:10.1016/j.tetlet.2005.02.124.

Further reading

External links