A wide variety of creatures inhabit the forests, tundras and bogs of the Lapland reserve. Each group of animals has its own unique role in a complex hierarchy of ecological relationships. Special place is hold by the most numerous predators in the microcosm – spiders. Оrder of spiders (Aranei) belongs to the class of the arachnids (Arachnida), subtype chelicerate arthropods (Chelicerata), type of arthropods (Arthropoda). At the present moment almost 44 000 species of spiders are known in the world. In number of terrestrial arthropods species, they are second only to related order of acarians (Acarina) and five major insect orders (beetles, butterflies, bugs, Diptera and Hymenoptera). Spiders have fully inhabited all natural areas and terranean habitat types. Spiders fauna in Russia has more than 2 500 species, and this is not the final number. It is not easy to present a complete portrait of these creatures; we will try to tell at least about several features of them. Unlike millipedes, crustaceans and insects spiders like all the chelicerate arthropods have quite simply organized mouthparts – there are two chelicerae (jaws analogue) and two pedipalps (tentaculites). Chelicerae consist of two segments – a main one and an end one, claw-like embedded in the main segment stria like a pen knife cutting edge. Clawlike segment by the way it is organized resembles a fang – inside is a channel which is used to conduct poison into a kill or an enemy body. However, chelicerae are used by spiders not only to bite an enemy, they also catch and mill kills; it is also a tool to clean the other extremities, dig soil, cut a web while constructing it and for other needs. Pedipalps unlike chelicerae Spiders of the Lapland reserve Spiders of the Lapland reserve 1 Lapland state nature biosphere reserve Spiders of the Lapland reserve consist of six segments with claws on the top of the latter and are mainly used as tactile organs, although this is not their single role. Using pedipalps spiders hold a kill and other objects, taste and smell, make a web. If fore feet are lost, pedipalps can partially replace them and help moving. Many species of spiders are able to cast feet. As a lizard’s tail, cast leg continues moving for a while, distracting potential enemies, and a new one grows on its place provided that a spider can still shed. Adult males’ pedipalps convert into a copulatory organ. The latter function of pedipalps is typical namely for spiders and is not encountered among the other chelicerate arthropods. Once upon a time ancestors of male spiders used pedipalps to transfer bags with semen – spermatophores – and passed them to females during a mating ritual. To make carrying spermatophores more convenient pedipalps have been changing gradually, a claw shifted from the top to the side surface of the last segment, and then acquired a bizarre shape, looking like a syringe capable of injecting semen into female genital pores. Segment itself has increased, acquired a spoon shape, and an onion-like syringe-claw securely storing sperm is located inside a “cup” of a “spoon”. The need for a protective bag – spermatophore – disappeared, and the only thing that reminds about it is that spider’s spermia are not located in semen one by one, but packed by several pieces together. Male pedipalps can often have some additional outgrowths, bristles and deepenings of various shapes that fit together female genitals key to a lock. This mechanism of “a lock and a key” is an obstacle for interspecific hybridization and therefore is an important generic feature. Drawings of the external sex organs are the basis for description of spider species in the scientific literature. Eyes of remote ancestors of spiders were faceted, complex, consisting of many simple eyes, just as of modern insects. However, after inhabiting ground ways of two groups of ancestor diverged – if insects preserved faceted eyes, individual facets of chelicerates were divided and gave start to formation of simple eyes. It should be noted that such eyes often give to an image of spider a remarkable resemblance to vertebrates, unattainable for other arthropods. Figure 8 is amusingly repeated manifold in a structure of spiders. This is known by everyone amount of walking legs, which distinguishes most arachnids; and a number of eyes (although some species of spiders, especially those living in caves can have reduced amount of eyes by one pair or more, until complete absence of eyes). Even body shape of a spider resembles an eight, consisting of the anterior part – prosoma and posterior – abdomen. Prosoma is a half of body that is responsible for interaction with the outside world. For a reason a head and sense receptor of bilateral animals with a front end of the body (as opposed to radially symmetrical animals, like sea stars) have developed namely in the front, in the direction of movement. This process reached its logical end at spiders: the front half of the body has not only main sense receptors, but limbs, and a nervous system inside a prosoma coordinates everything, it is concentrated in a rather complicated for invertebrate brain, more perfect than, say, insects. Even di by m no a The pair of spiders Microlinyphia pusilla: differences between pedipalps of male and female are well seen Look of the spider-wolf Alopecosa aculeata seems quite sensible The spider-wolf Trochosa spinipalpis, side-view 2 ABCGheritage – Our common arctic heritage al A fa ov ar in un in an O fro po or to en le m pe m ne th It of ev th sw ge th er in te bo le to of tia er 4 of Figure 8 is amusingly repeated manifold in a structure of spiders. This is amount of walking legs; a number of eyes and even body shape of a spider resembles an eight. We will again remember extinct predecessors of spiders. On rear part of a body they had limbs that were inherited from the more ancient progenitors of all arthropods – annelid polychaete worms. It must be said that limbs of animals – are organs than can in the course of evolution be easily subjected to amazing modifications and can switch to completely different roles than in the beginning. This happened to abdominal legs of ancient chelicerates: a part of them, which served for movement and breathing in water (they settled gill branches) persistently kept on perform a respiratory function when animals began to inhabit ground. For this purpose, pits and later nests for respiratory legs were formed in abdomen to protect them from drying in air. Thus limbs transformed into lungs. It distinguishes spiders from insects and strangely reminiscent of terrestrial vertebrates: only they and we have lungs. However, with decrease in body size insect respiratory system – the trachea – is more suitable and many groups of spiders switched to it partially or completely, and very small species generally lost their respiratory organs and breathe through the whole body surface. Hind abdominal legs of ancient pre-spiders fulfilled another role. Spinning glands appeared namely here. What was their initial function is difficult to say; possible for making a shelter or protective canopy for eggs laying. Over time, the main body of the glands moved to abdomen, only glands outputs left on shortening legs; number of legs segments was reduced to two or three, and the legs themselves displaced to the end of abdomen. Thus spinnerets were formed, by the way an initial number of them was equal to eight (four pairs)! However, amount of spinnerets of modern spiders reduced up to 6, 4 or 2; it is enough to create webs. One pair of spinnerets of some spiders species is still very elongated and resembles small legs, often used in addition to creating web as sense of touch. Position of spinnerets on the end of the body, even a hind one, has proved to be very successful: they are able to move as hand fingers, abdomen can also be moved from side to side, moving the entire “palm with fingers” like a painter’s brush. Amount of waking legs appeared to be abundant, therefore a rear pair is used, in addition to walking, to pull a web and make manipulations with it. So it is not a surprise that with such a set of “tools: and freedom of movement spiders have developed and improved their web making abilities. Suitable body structure served as a basis for further evolution. Prosoma and abdomen of spiders are connected with a small pedicle which connects all body parts – blood vessels, nerves and intestines. And that’s basically all that is contained in the pedicle! It is very thin, so it can be used as a special “joint” for abdomen movement, at that abdomen itself can not change its shape and operate easily, being more mobile than a prosoma. On the other hand, such a narrow pedicle testifies about a perfect internal structure of spiders allowing connecting the fore and hind body parts by minimum amount of “binding yarns.” The whole spider body is covered with hairs and setae being more important than it may seem at the first glance. The fact that unlike us, having soft skin which is direct contact with the outside environment, arthropods’ body is covered by a protective carapace – cuticle, which outer layer almost doesn’t content living cells. There are advantages and disadvantages. For example, a cuticle can be attached muscles and another organs from inside, and a body retains moisture for a long time thanks to such a carapace. And on the contrary, it is impossible to grow within it, so spiders have to shed from time to time, getting rid of an old “fell”, becoming softer and defenseless, and body size significantly increases only at that period. Hard cover restricts mobility, and body with limbs have to be divided into segments connected by more elastic parts (hence the name of species is arthropods). We, chordates, were also originated from annelids, but went the other way – created an internal skeleton and preserved elastic alive external covers. Each way was good: arthropods successfully occupied a niche of small animals and chordates – medium and large. So a protective hard cuticle makes it almost impossible to feel anything. And the most part of sensations – tactile, chemical (smell and taste), auditory is transferred through a cuticle from different types of hairs as if dressing a rigid spider’s body in the second ephemeral sensitive shell. In addition, hairs protect a body from contact and water (the bigger is a spider, the thicker hair covering it has, the more it resembles wool of mammals), create on its hind legs series for spinning a web and series in sound bodies, thicken into spikes for catching preys on the front legs and form tenacious pads allowing walking on any surface ... and finally, hairs are mainly responsible for spiders coloring. Color of an animal is determined either by presence of pigments in tissues or structural features of the latter. For example, white spots on abdomen of diadem spiders (genus Araneus) are formed by deposition of guanine, and metallic sheen on Spiders of the Lapland reserve digestive system in spider’s prosoma is represented primarily by departments responsible for interaction with the environment – esophagus and sucking stomach sucking (spiders do not swallow food using jaws and throat, but gradually suck it in a liquid form using a “vacuum pump” – sucking stomach). The rear half of spider’s body, by contrast, has collected almost all organs responsible for internal processes in a body. A considerable part of the digestive system, and a chemical factory: – liver, excretory system, reproductive glands where ova and sperm are formed are located here; spinning gland are located here as well, capable of synthesizing unsurpassed in its quality protein building material. If the main in a prosoma uniting other organ systems (limbs, senses, etc.) – is a brain, in abdomen such a center is heart ensuring hemolymph run and metabolism. 3 Lapland state nature biosphere reserve Spiders of the Lapland reserve Xerolycosa nemoralis – one of the most brightly colored spiders-wolves in reserve Abdomen of Araneus quadratus has white spots forming quadrangle body of leaping spiders (Heliophanus) is manifested by refraction of light in lenticular areas in cover hairs. Coloring plays a protective role, allowing a spider to blend with a surrounding substrate – the so-called cryptic coloration. Spots and stripes can conceal an overall silhouette, in this case coloring is disruptive. Sometimes a body shape and color of spider resemble other creatures – mostly ants – it is mimicry. All mentioned variations have one main purpose – protection from birds, because namely birds among the other predators that eat spiders, are mostly oriented on vision while hunting a prey. Sometimes coloration may be physiological – match the original color of tissues and body cover. Some species are darker than the others and thus heat up quickly in sunlight, spiders sitting on their webs “back down”, have a dark color on the underside of a body, usually facing the sky. Coloration of spiders with good eyesight also has a signal role. Origin of a pattern on top of abdomen is interesting – it is often presented by a tapered longitudinal spot in the front half and wide “herringbone” in the rear part. So, a tapered spot is usually located just above spider’s heart and copies its size and shape. Perhaps it serves as heart temperature regulation, as differs in tone from the rest of abdomen coloration. And a tapering “herringbone” of repetitive elements – this is the only externally visible trace of posterior half of a body segments connection, which once led to formation of abdomen. It should be noted that segmentation of chelicerates body is determined by genetic mechanisms similar to those of vertebrates (certainly vertebrates’ body is not segmented externally, but the skeleton has repeated segments, such as vertebrae) and it is coded differently at insects comparing to us and spiders. If you look at the overall body structure, its uniformity within the order is striking, no step aside. The same can be said about feed preferences: all spiders, with the minor exception – are certainly predators (only some species can eat pollen of plants, one lives on protein bodies of tropical acacia intended for ant-trees symbionts). Heliophanus dubius male glitters brightly in the sun These two features are typical for the other groups of arachnids, except acarians. These are namely spiders and acarians which reached an unprecedented prosperity among the other chelicerates. Acarians have achieved this by switching to a variety of feed sources, and due to their ability to change body structure with the course of the evolution; it means due to a very deep and multidirectional specialization. Spiders, on the contrary, remain omnivorous predators and the structure of their bodies is the least specialized of all orders of arachnids. The set of their limbs is slightly abundant, therefore legs easily acquire an additional role in case of minimal structure changes and are used to hold prey, feel a substrate, for manipulation during web making, some tropical species – for throwing spider “lasso” with a drop of glue on the end or catching insects with an elastic “butterfly net”, for fixing a female during copulation, etc. Loss of any leg or pedipalp is easily compensated functionally by adjacent limbs. Therefore, lifestyle changes for spiders became widely possible without a fundamental restructuring of body. But other orders of arachnids, by contrast, even at the moment of appearance followed the way of narrow morphological specialization. Let us consider, for example, scorpions. Their pedipalps turned into claws, abdomen has a stretched section-“tail” with a poisonous needle on the end, thin walking legs located in one plane are unable to support body by weight – scorpions were not able to retreat the structure formed in the aquatic environment. It defines possible sizes, and lifestyles, imposing strict limits on diversity and limiting the number of emerging species. On the other hand, non-specialized common body structure of spiders is complimented by truly fantastic opportunities for using a web. Different taxons, little changed outwardly, mastered new variants of hunting spider facilities and took new ecological niches. Even aquatic spiders surprisingly little changed comparing to terrestrial spiders, so their appearance cannot be used to identify their habits; but one glance on sub- 4 ABCGheritage – Our common arctic heritage m fo fib co in th up po an fla sh en gl ar a to nrs ren w e, as d, rt cle m- cniy, ok le ce b- marine bells created by them and filled with water is enough for it. Technical characteristics of a web transcend any man-made fibers, that clearly indicates perfection of spider silk chemical composition, acquired as a result of the evolution. Depending on the purpose the same spider can create multiple spider threads of different composition and different texture, using up to seven types of silk glands. So, diadem spiders using ampoule-like glands secrete a thick dry dragline of a radial network and a thin safety thread when moving outside a web; tree and flagelliform glands secrete sticky spiral thread of a web: pearshaped glands form bundles of thin adhesive fibers, used to fix ends of a thick filaments to objects; threads of segment-like glands are used to mantle prey; threads of tube-like glands are used to form an egg cocoon, etc. When building a web a spider carefully calibrates thread tension and acts according to a complex instinctive action pattern. At that a catching web Arctosa alpigena – spider-wolf with “a white heart” can be modified depending on structural features of a place where it is being built, weather conditions and other factors, and sometimes can be modified significantly. Thus, orb-web spiders in certain conditions may not make a full web, but end with a small disorderly weaved threads, for example in case of prey abundance – for example, in a street lamp light. There is an original way to define some toxic substances – liquid with admixture of suspicious matter is given to a spider to drink and then they wait until spider makes a capturing web, change of its shape is used to determine exactly what kind toxin has been detected. At the same time webs of orb-web spiders have such characteristics, that they can be used to identify species of a host, and some of these features (constant ratio between the sizes of the individual parts of the network) is on the networks of this spider species at any age. Except building catching nets, web is used for moving safety (“trace thread”) for mantling a prey and preparing space for Spiders of the Lapland reserve hns hto ge ue rs, crs eal e, – or e- Alopecosa aculeata – spider-wolf with “a dark heart” Araneus marmoreus extracts an insect caught into his web Male and female Nuctenea silvicultrix 5 Lapland state nature biosphere reserve Spiders of the Lapland reserve its storage and eating, small species and young spiders of large species – for building shelters, threading burrows, weaving egg cocoons, for construction of spermatic net by males (which they use to fill their pedipalps with a seminal fluid from abdomen), to mark territory and leaving odorous marks, hanging while shedding, to create air domes by water spiders. Another direction of spiders’ biochemical evolution, where they also achieved perfection – is production of poison. Spider’s poison consists of mixture of various biologically active substances – from salt to high molecular proteins – and includes more than a hundred of components, which is a record among injurious animals. This complex of substances works in alignment, providing high efficiency of total action; poison composition is specific for each species of spiders and may also depend on age, sex, feed, etc. In most cases effect of poison is directed at main preys of spiders – insects (at that specific toxins may pose a risk only to a very narrow circle of preys); mechanism of action is neurotoxic or necrotic. Poisons are often intended also for protection from enemies, therefore there species of spider dangerous for humans – composition of their poison is configured to repel vertebrates. In our country, such spiders inhabit southern semi-desert and called black widows. Diversity of spiders species makes their poisons inexhaustible and irreplaceable natural biochemical card index of active substances used in neurobiological research and medicine. The question arises: why spiders species fauna is so rich even in the northern territories such as the Lapland reserve? Spiders almost do not specialize in different feed sources, like many other animals (for example, snakes of the central Russia: viper hunts small animals, water snake – mainly amphibians, smooth snake – lizards). Hence, separation of ecological niches of spiders occur on other grounds. First of all this is a different feeding behavior (which includes use of different types of catching nets) due to which one habitat can be used differently. Let us review, for example, a pine forest of the reserve near Chunozero settlement. Between neighboring trees an orb-web spider Nuctenea silvicultrix is weaving its web, a diadem spider Araneus marmoreus makes its webs on main pine branches, horizontal webs of long-legged Bolephthyphantes index appear in thickets of small branches, small fine hammocks of Obscuriphantes obscures are seen among large needles; simple nets of marked Drapetisca socialis are stretched directly over cortex, cavity under cortex is densely filled with hunting nets of Theridion varians, microscopic threads of tiny Savignia frontata are almost invisible in truck bottom cavities. Species hunting without nets are inhabiting near artificial air threads – slow flat crab spiders Coriarachne depressa and gracious mobile crab spiders Philodromus fuscomarginatus live in splits, dark-gray twilight hunters Gnaphosa lapponum spend their time under wide layers of dead bark, bright striped Salticus cingulatus hunt their preys on a sunny side of a trunk, big colored spiders-wolves Acantholycosa lignaria keep territory near roots. These are the most common inhabitants of the pines. Firs and birches are inhabited by their own species… Thus spiders can live and hunt on a bark of pine in several different ways, and the same method can be accustomed Male Bolephthyphantes index The web of Obscuriphantes obscurus Coriarachne depressa 6 ABCGheritage – Our common arctic heritage After fog dew allows to see subtle travel threads of spiders Eating a fly the female Bolephthyphantes index is a feed for a larvae of parasitic fly at the same time Spiders of the Lapland reserve Salticus cingulatus to different substrates – bark, stones, ground, and finally spiders are turned out to be dependent on substrates as well as a method of hunting. So, different methods of hunting, multiplied by structurally different substrates – is one of the main reasons explaining coexistence of a large number of spiders species at the same habitat. Spider population can be very large and close to the number of all the insects of the same size in a given area. If you are lucky, in a quiet autumn morning when dew is falling or after a small rain you can see it with your eyes, when usually invisible nets are sparkling silver water droplets systems, and even one step can disturb at least one of them. Amount of small spiders in litter fall per 1 m2 can reach more than 200 zooids, and population of one tree – a few hundred of representatives from tens of species. It seems that it may not be enough prey for such hordes of predators and seeming abundance of spiders was noted by researchers more than once. It is explained by several reasons. First of all, spiders fully digest feed (general ecological rule, stating that only 10 % of ingested biomass transfers to the next level of a food chain is not applicable here); Secondly, the vast majority of spiders is capable to live without feed for a long time not harming themselves; Thirdly being non-specialized predators when other feed is absent they can eat their own kind. Therefore biomass of spiders is safe with respect to adverse and changing environmental conditions, and their population from year to year remains very stable. It is clear that the role of such persistent and massive predators in regulating the number of insects is enormous. A main enemy of spiders are birds, and in they can be a basic ration for many insectivorous species such as tits. Shrews, viviparous lizards, ants, predatory beetles and many other creatures also readily consume eight-legged hunter. At the moment, the Lapland reserve is inhabited by more than 270 known species of spiders belonging to 17 families. They are found for 4 years of researches, and the fauna turned to be rich enough for a relatively small northern area. The reason for this abundance lies in diversity of habitats in the reserve, and a significant contribution for this is made by hills – Chunatundra, Monchetundra and Salnye tundras. Their hillsides are open to different sides of the world and are illuminated differently by sun; each elevation is characterized by its vegetation; rock yields in form of boulder streams, cliffs and rock fall are found on the slopes. All these conditions create a rich mosaic of habitats. Tundra also insulates well watersheds of rivers, which help different species to live in similar but distant locations. By hunting strategies spiders are divided into webspinner spiders and hunters which do not use trapping nets. More differences are found within these groups. Looking at appearance of a spider one can immediately determine its typical way of obtaining feed, so its external structure is closely associated with feeding behavior. Active hunting without using a web is considered to be an initial strategy, however, some modern hunters have webspinners as their ancestors. 7 Lapland state nature biosphere reserve Young Dolomedes fimbriatus The marshland on the lake Spiders of the Lapland reserve Close families of spiders-wolves (Lycosiae) and spiders-ramblers (Pisauridae) belong to the group of daytime hunters. They have a rather large size, long strong legs, well-developed eyes and protective coloring in the color of ground, stones, or leaves; they run and jump well. Most of these spiders spend life in constant motion and therefore often found underfoot in the woods and swamps. Inaction is alien to them, except only at the onset of cold weather; active lifestyle requires constant feeding, therefore feed absence is more complicated for these spiders comparing with another. Dolomedes fimbriatus is the one representative of Pisauridae in the reserve – the largest species among terrestrial species, reaches more than 4 cm in legs span. Inhabits grassy marshes and coasts, sometimes hunts even aquatic animals. In north conditions for large species of spiders it is very important to choose a warm habitat and swamps in this regard are very suitable biotopes for size record-breaking spiders of the local fauna – protected by forest from the, exposed to sun, storing heat in wet moss cushion for a long time. A life cycle of this spider in the reserve is two years, mature spiders appear in June, after the last shedding. Many species of spiders when mating period begins in early warm season (May-June), cannot shed for the last time and become sexually mature, not wintered. Spiders overwinter in leaf litters and can completely freeze without any threat to life due to natural antifreeze contained in hemolymph. Throughout the summer and early autumn spiders of the penultimate stage of development accumulate fed and only hibernation stimulates them to shed – this is the mechanism of simultaneous mass emergence of adults in early summer. Male finds a female spider, approaches and performs a courtship ritual. Often it is waiting for a suitable moment when a female catches a prey and begins to eat – it is safer. After mating, a female feeds up and constructs a large spherical cocoon in June that is always carried by her with some difficulties. In some time it chooses a place and hangs a cocoon on sedge leaves or heather twig, and then it spins around a large loose shield ball, of a grapefruit size of strong messy threads and remains in this ball to protect a cocoon. It repels in an of re th ly or fro on of fir to ca sm es by Th Pr to on an tio un an do w N ce a ve fe of an (s on Growth stages of dolomedes: kids in the nest, grown up spider before its first hibernation, young female before its second hibernation, adult female 8 ABCGheritage – Our common arctic heritage ms nt er. rme ond sy els insects by itself, rushing at them and hitting with front legs, and a ball is serving as protection from birds. Several hundreds of baby spiders leave a cocoon in August, for 2–3 days they remain as a single bunch inside a ball, and then scatter around the area. Typical light stria appear at baby spiders pretty early, but the overall coloration is not velvety brown, but olive or green; they live in grass stand, often encountered quite far from adults. In September baby spiders of the first and second years of life may acquire a reddish outfit to match colors of autumn leaves. In calm sunny autumn days after cold nights babies of the first year are spread through a flight. To do this, they climb on top of a high plant, stand on tiptoe and raise abdomen vertically, taking the starting position. Sun-heated air flows them smoothly up and at this moment spinnerets start evolving finest strong thread. The thread is getting longer and picked up by the air at some point almost takes a spider from surface. Then it attaches a thread end next to it, moves to it and bites. Probably all kinds of spiders found in the reserve are capable to fly. Autumn journeys in search of new territories are limited only by aeronauts size, so only baby spiders of large species and adults as well of small species are flying. Apart from the main life cycle – growth and reproduction – spiders and dependent on shedding cycles. Dolomedes undergoes throughout life about 8–10 sheds until it becomes an adult. During shedding a spider is absolutely helpless and does not respond to external stimuli, fully concentrating on what is happening. A usual way to shed is hanging back down. Number of external and internal factors can disrupt this process and a spider can lose a limb or die. Shedding occurs in a hidden place, often at night. Immediately after it covers are very elastic, and a spider is unable to move normally and defend itself, it easily loses and absorbs moisture. After a couple of hours a cuticle hardens, coloration becomes brighter and another inter-shedding period begins. Relatives of spiders (scorpions, amblypygi), and many tropical spiders (though only female) can shed being adults unlimited number of times. At the moment the Lapland reserve is inhabited by more than 270 known species of spiders belonging to 17 families. The grass spider-crab before flying Spiders of the Lapland reserve re es on alnd al er elm r- The female under the dome of the ball Balloon-nest 9 Lapland state nature biosphere reserve Pardosa lugubris: immature May male, adult June, and July female with cocoon amentata, dark Pardosa paludicola and small Pardosa prativaga) to the highest tundra peaks (Pardosa eiseni inhabiting stony areas and Pardosa palustris of a color of dried grass). The first two species are common in the moderate climate and their breeding season there starts very early – in April. Here in the north, along with the other species they start breeding in summer. One species from the top of Salnye tundras Pardosa indecora, should not be found on the Kola peninsula at all, as according to all data it is encountered far to the east, for example, on Taimyr. Representatives of the genus Acantholycosa changed free lifestyle to more sedentary. Close relatives of Pardosa, they differ by large size and long legs. Each female owns its personal territory of several large stones (acantholycosa norvegica) or a trunk of a fallen tree (acantholycosa A. lignaria), where a spider hides in case of danger. If one cautiously approaches an area inhabited by acantholycosa spiders, he/she can make good pictures of sitting spiders, observe them hunting. Among Lycosidae spiders there are also amphibiotic spiders living along banks, marshes, craftily walking by water and Spiders of the Lapland reserve So why are our species not capable for that? The main reason – hard external sex organs – a special plate with holes on abdomen of a female and modified pedipalps claws of a male. Beneath new covers are not able to be formed, and shedding would lead to tearing of tissues. Thus, any adult male spiders are not capable of shedding, but female of species without abdominal plates can usually do it and live longer. Generally, such species are spread in countries with a warm climate. The largest Lycosidae of the reserve Alopecosa pinetorum together with a short-legged phlegmatic Trochosa spinipalpis, like a fishing spider is encountered on margins of cozy and warm swamps and is very rare. More common is a smaller Alopecosa aculeata – the most notable and active spider on a dry fragrant needle bed of sparse coniferous forests in June. The majority of Lycosidae is represented by Pardosa, agile and graceful small hunters, each species of them prefers a different type of soil and a plant litter. Thus, all 13 species of Pardosa share the territory of the reserve, although often representatives of 2–3 species can be found together. They live everywhere, from marshes and shores (colored Pardosa Common in tundra Pardosa eiseni is basking in the sun Pardosa paludicola 10 ABCGheritage – Our common arctic heritage a) rwo dh, er. a, de, ee ey na) re es ke pind A. norvegica Female Pirata piraticus Spiders of the Lapland reserve Acantholycosa lignaria diving if necessary. In the Lapland reserve such species of spiders from Pirata genus are represented. They have a slightly greenish color with a light trident on cephalothorax. The most common is Pirata piraticus, and its close relative, actually reduced copy of it, Pirata insularis, is met only on a sedge swamp around a steam at the bottom of the ecological path “Elnyun-II”. Cocoon of Lycosidae is smaller than the one of Dolomedes. Usually females attach it to spinnerets and quietly go about their business, not forgetting to use every opportunity to sit in the sun, warming their future offspring. After hatching spiders remain protected in a swollen and shed there twice before to go outside. Baby spiders immediately get on a mother’s body and hold its hair. At certain times of a year Lycosidae females covered by baies can be found everywhere. This characteristic period for the family of protecting offspring lasts several days, during the period a mother is gradually throwing off baby spiders in different places it likes. Maternal affairs do not prevent it from regular hunt, and, if weather allows, the first cocoon will be replaced by the second one and occasionally it manages to get the third one. A peak of evolution among day stray spiders – are jumping spiders (family Salticidae). Their world perception is based on vision, like human. Central front eyes are very large and are not spherical but tube-shaped resembling a pair of binoculars, which they are in fact. They really do give a detailed enlarged image. “Binoculars” field of vision is small, and this defect is compensated by two mechanisms: main eyes are able to move and follow an object, and provide all-round view is ensured by remaining 6 eyes. The front side eyes fall short of size comparing to central ones. Their function is different: they are not responsible for enlargement, but for binocular vision (three-dimensional perception of space) as they are located most widely apart. Other eyes give a low quality image and serve to expand a visual field. This function is performed by rear eyes of all spiders, but it is perfect at jumping spiders – only their rare side pair is pushed back, sometimes behind the middle of prosoma. Generally speaking, it is difficult to imagine how jumping spider see the world: to usual for us color vision they have additional built-in binoculars and back view eyes and it all works at the same time! Vision allowed jumping spiders to master the technique of jumping, which they use to attack (short rushes creeping up a prey at the right distance) and movement. These capabilities are particularly useful when moving in the midst of plants: looking all around and accurately estimating a distance between objects, the spiders jump between leaves and stems, as they want. Movement in a complex three-dimensional environment requires development of a nervous system, and, like other animal having similar lifestyle (primates, arboreal amphibians, parrots), jumping spiders are the “smartest” among their relatives, in this case – among spiders. It is proved that learning by trial and error can be a mandatory component of jumping spiders behavior unlike fully programmed instinctive actions of most other spiders. 11 Lapland state nature biosphere reserve Female Acantholycosa norvegica with cocoon Female Pardosa prativaga with cocoon Many jumping spiders acquired bright coloration, because they are able to see and appreciate it. Dresses of all rainbow colors are used to recognize zooids of their own species, to distinguish males, females and baby spiders, for refined mating dances. Jumping spiders are usually quite small, but when looking closer to these living colorful sparks, few can resist the fascination. Thick body with compact limbs covered with thick hair, and “owl-like” expressive eyes changed a standard spider appearance beyond recognition, giving the jumping spiders the appearance of mysterious animals. Getting accustomed to something, they turn cephalothorax following an object producing a lasting impression of an “intelligent” view. The Salticidae are abundant in the tropics, where they can even prevail in number of species among the rest of the spiders. This is the largest family of spiders in the world consisting of more than five and a half thousand species. To the north of the equator, their diversity is steadily declining, and the fauna of the Lapland reserve is represented only by 7 species of jumping spiders. The greatest number of these thermophilic spiders met is encountered in the western slope of the tract Saidnotlag, in a rocky hollow, closed from all sides except the south. Good drainage at constant streams fall, heating by direct sun- light and protection from northern winds create there a very special microclimate. This unique area is inhabited by a number of southern species not encountered anywhere else in the reserve. We can assume that such emigrees are periodically brought here by southerly winds; flying over Chunozero lake and coming across Chuna tundra mountain slope air waves leave tens of spiders of different species on a warm slope. In addition to daytime hunters, there are twilight and night hunters, having an elongated body and almost equal-sized eyes, usually located in two rows. Close families of sac spiders (Clubionidae) belong to them: Zoridae, Liocranidae, as well as Gnaphosidae. Unlike wolf spider running and jumping everywhere, ground spiders hyenas are secret ramblers with a monotone color and white, as if lifeless eyes. This color is due to a silvery reflective mirror, typical for many nocturnal creatures. However, a part of eye remains dark, day part. Most spiders have mirrored eyes, but eyes of Gnaphosidae are especially noticeable. Representatives of different genera and species are remarkably similar, only size and gloomy shade colors vary. During a day they hide under bark and stones, they also overwinter there and protect cocoons attached to plane substrates. th ha sm of by Th by se in Spiders of the Lapland reserve ad on in sp m su Young female Evarcha arcuata Immature female Evarcha falcata Immature male Evarcha falcata 12 ABCGheritage – Our common arctic heritage ca to bl to qu Adult female Evarcha falcata Adult male Evarcha falcata ery ber relly nd ave ht es, lunaFemale Sitticus saxicola Only two species of Micaria genus sharply distinguish from the total mass which switched to a day lifestyle. However, they have a capping relation with other ground spiders. Well looked small micaria run in a day time copying habits and appearance of small ants and wasps. Their dark body sparkles in the sun by all rainbow colors as it is covered by hairs refracting light. The two most common species of this genus divided simply by their preferences – one lives in the lowland part of the reserve, and the other – on the hills; other species are met only in Saidnotlag mentioned above. All males from the Liocranidae and sac spiders, including all adult females from the first family (genus Agroeca) were found only in August and September, i.e. their mating season starts in the very end of the warm season. For medium and large spiders of the Lapland reserve it is an exception to the rule – males of other species of this size group mature in early to mid summer. Among spiders-hunters not making webs spiders-ambuscaders stand alone – “living traps”. If wolf spiders are adapted to rush, jumping spider – to jump, spiders hyenas – to “ramble”, a business card of ambuscaders is their agility, ability to crab movement, turn quickly to sides. Such skills were required for inhabiting limited space, such as a fracture under Zora spinimana Spiders of the Lapland reserve nd nd ve art es, enar, de ect Slope of the tract Saidnotlag 13 Lapland state nature biosphere reserve Spiders of the Lapland reserve Female Gnaphosa Up: Agroeca proxima, down: Clubiona Immature male Gnaphosa Worried female Gnaphosa tree bark, a flower, a cavity between soil particles. Maneuverability is achieved very simply: by a small rotation of an axis attaching legs to a body, as a result legs are not only bent downwards, but sideways as well. System of spider’s legs reminds a set of tongs, a front pair is bent forward, and hind legs – often – backwards. Two close families of crabs – ambuscaders (called spider-crabs because of an appearance and manner of movement), Philodromidae and Thomisidae differ in a relative size of front and rear pairs of legs – legs of the first are about the same, and rear legs of the second one are shorter than front pairs. Front legs are equipped with numerous thick spikes, helping to keep even a very large prey. Eyes are only day, they are directed to different sides and raised above the surface of prosoma for better viewing, like other ambuscaders – frogs, chameleons, flounder, mantis. Cryptic coloration of crab spiders is adapted to specific conditions and is much better than, say, coloration of wolf spiders, which has a generalized character, more or less suitable for other habitats. Floral spider Misumena vatia, found near Lower Chuna, is even capable of changing a color of its body, like chameleons, but slower. Its arsenal consists of green, yellow and white dresses. Philodromus fuscomarginatus’ color tone is also changeable, it inhabits pines, spiders of a red and gray colors are encountered. Body of crab spiders inhabiting tree bark is often flattened which only increases the similarity with some crabs hiding among the rocks on the bottom. All Philodromidae are featured by high speed and maneuverability, but as in other families, each genus prefers a different type of substrate: Philodromus – bark, Thanatus – soil surface, Tibellus – grass. Grass – a special place to live, and a body of tibellus is stretched, covered with long longitudinal stripes; stopped dead in ambuscade missed – a spider looks like a straw. Thomisidae on the contrary are slow (but “agile”). Besides mentioned Coriarachne and flower spiders they include numerous gray-brown “crabs” of the genus Xysticus (7 species) that live on ground and low plants, and several small species of Ozyptila, similar to baby Xysticus. Diminutiveness of Ozyp- 14 ABCGheritage – Our common arctic heritage til m cl co ar da se bo pa a ufoil nd al ks es us) es p- Misumena vatia on Dwarf cornel tila allows them to penetrate deep into a plant litter, shrubs, moss cushion. Thick hairs on their bodies partially became claviform – such tiny upraised “cams” that protect a body from coarse grains. A main enemy of spiders are birds. Shrews, viviparous lizards, ants, predatory beetles and many other creatures also can consume the spider. Among nearly invisible inhabitants of reserved bed there are spiders with very primitive prey networks (family Hahnidae). Web apparatus of most of Theridiidae consists of 6 long segments spinnerets located strictly in a row across abdomen bottom. Using spinnerets spiders construct a horizontal flat panel of irregular shape and go over the network. In fact, such a network – is just a better copy of soil surface, where prey can Spider-ambuscader in hunting pose stick, thread tension allows to understand location of a kill. Externally spiders of genera Hahnia and Antistea resemble small gray night hunters than “real” Theridiidae. Only Cryphoeca silvicola, living in birch bark is more or less brightly colored. Its web apparatus is not exotically structured. But this species does not avoid a litter – in autumn cryphoeca go down there for hibernation (crab spiders Philodromus do not do it and remain for overwinter under bark). On the contrary makers of perfect wheel-shaped vertical webs are not encountered in a litter. These spiders inhabit all sorts of vegetation and rock slides. Their webs consist of radial threads forming a kind of bicycle spokes and spiral sticky trapping thread stretched between them. Such an arrangement allows catching an impressive space sector with minimum material consumption and repairing damages easily (as tension of each section of a network can be adjusted separately). Working surface is attached to a simple frame, usually triangular, only three points of support are enough to fix it; thick threads of a random length drawn on these points. Finally a network can be placed practically everywhere. Usual- Spiders of the Lapland reserve rtnds frs of ve ut an es, ey of gs, pin, cugal ses, ab nks Color options of female Philodromus fuscomarginatus 15 Lapland state nature biosphere reserve Closely species of Tibellus – maritimus and oblongus Cryphorca silvicola Ground crab Xysticus audax has perfect protective coloration ly orb-web spider (Araneidae) construct webs at night and in morning when air is quiet, and majority of birds is sleeping. In soft light of a cool night busy silent constructors scurry between branches, sit in a center of networks, patch holes made by large moths, mantle preys and carry them into shelter near a network. Many species build a new web every night which is suitable for that and leave an old one, while others are able to collect used webs with small dipterans in them and eat a lump. Some features of radial networks and their creators have been already mentioned above. One can tell much about them; detailed descriptions are found in many books devoted to spiders. “Economic” effect of using radial webs is manifested by the fact that the largest Theridiidae of the reserve belong to orb-web spiders. Large terrestrial spiders are in better conditions – soil surface warms up well during a day, and hunters may find vitally required heat. Theridiidae living on air are harder to do it. tim of an N th m pe am is su la of on fro Spiders of the Lapland reserve To the left: female Xysticus ulmi, to the right: female Xysticus obscurus Ozyptila atomaria Araneus marmoreus on ground 16 ABCGheritage – Our common arctic heritage A pair of Larinioides patagiatus However, some orb-web spider find a way out – during day time they come down to a warm bedding, and representatives of one species – Nuctenea silvicultrix sit under pine trees bark and digest feed consumed in night time warmed by sunshine. Not for nothing large zooids of this species are encountered throughout a season, they can be found immediately after melting of snow. In more southern latitudes during breeding period of different orb-web species is spaced in time. For example, for a genus Larinioides mating season in the midland is typical in spring, for many diadem spiders (Araneus) – in late summer, for Araniella – in early summer; but within the Lapland reserve all of them have to move these important stages of breeding cycles to the hottest time – mid-summer. Diadem spiders often have a complex pattern of spots on an abdomen, and this pattern distinguishes the species from each other. Moreover, Araneus marmoreus has sev- eral options for coloration – gray, yellow and reddish. This marking serves for the same goal – protection, it conceals a silhouette of immovable spider. In case of evident danger diadem spiders just fall off a network and freeze. Large globular abdomen and thin legs of the Theridiidae simply do not allow them to escape from danger, and diadem spiders rely on their assumed non-visibility. Nuctenea is painted in gray tones of a bark color, Araniella cucurbitina living in shrubs – is of a clean green color, Araneus saevus amazingly merges with surface of dark stones on rockslides. Among small species of orb-web spiders only Cercidia prominens is common, it has a hard shell on abdomen, protecting it from predators; species of the genus Hypsosinga and Cyclosa are rare. Much more elegant are some species close to orb-web spiders from a small family Tetragnathidae. A common feature Spiders of the Lapland reserve nd g. ede ar ch le at rs ut ed tng ntre Immature females Araneus marmoreus have similar coloring (first photo), but adult females have different one 17 Lapland state nature biosphere reserve Spiders of the Lapland reserve Female Araneus saevus of this family – is elongated lobe-shaped outgrowths of initial pedipalp segment forming a kind of the second jaws under chelicerae (name Tetragnatha is translated as “four jaws”). Such blades, but shorter, are typical to the other spiders as well; they limit a perioral area. Structure of a radial network also differs – at Tetragnathidae its center is empty, at orb-web spiders it is usually tightened by a little messy mesh. Metellina merianae is so similar to the genus of diadem spiders, that scientists-taxonomists sometimes make it “jump” from one family to another. This species lives in caves, boulder stream, basements, etc., preferring a mountain landscape. Metellina is so dependent on its habitat that lives in appropriate places everywhere almost regardless of latitude. It can be found in the mountains of Central Asia and in the Lapland reserve, in the Carpathians and the Urals. Metellina’s web is asymmetric, its lower half has two times more semicircular trapping threads than an upper one. The genus Tetragnatha giving its name to the family includes only slim long-legged spiders which elongated body shape means adaptation to live in grass. Small neat meshes of elongated Tetragnatha Tetragnatha extensa, described by Carl Linne, can be seen on marshes, along river and lake coasts, among different grasses of the reserve. Adult spiders are encountered in summer, and baby spiders can be found at any time; in autumn growing spiders often become reddish, similar to growing Dolomedes spiders. The smallest have almost spherical abdomen, which is increasingly stretched while growing. When hunting Tetragnatha are stretched out in a center of their networks and externally resemble a needle. In case of danger they run across to a stem or leaf of grass next to a network, embrace it with shortened legs of the third pairs and spread out to become almost invisible. As hard to imagine riverbanks without sedges, so hard is to imagine riverbank sedge without Tetragnatha. Female Cyclosa conica Th of na th on Ve co am is sim to is ro la Metellina female A pair Tetragnatha 18 ABCGheritage – Our common arctic heritage re fro of w liv ar lo da sp ho at in ha ca w in fu a bi in its Orb-web spiders are on the top of the evolution among the Theridiidae, as jumping spiders among hunters. However, ways of evolution are fancy, and the last from two genera Tetragnathidae – Pachygnatha found in the reserve includes species that do not use trapping networks at all and returned to live on simple hunting. Pachygnatha clercki is found only in the entry of the river Verkhnyaya Chuna; spider roams on a surface of soil across coastal vegetation, using transportation threads to move among sedge stems sticking out of water, and if there is a need – it can run fast. Externally pachygnata is not really similar to its relatives Tetragnatha and Metellina but similar to them is widely spread, from Central Asia to Taimyr, which is associated with its choice of habitats – stone deposits and rocks, water coasts and sedges can be found in almost any latitude and longitude. The only predators that could permanently settle in human residential buildings also belong to the Theridiidae, primarily from the family of the cobweb spiders (Theridiidae). Species of Steadoda genus, distant relatives of black widow, are worldwide common in residential buildings. In the wild nature they live in dry rocks and caves of warm countries, therefore they are capable of staying without feed and water for extremely long period, and their web network functions well for many days with a minimum repair requirement. Real synanthropic spiders are not encountered in the reserve’s farm – as internal house temperature in winter is too unstable; but a local steatoda species small Steatoda bipunctata inhabits the surrounding territory, in the woods, under tree bark. It is eager to inhabit the houses inside and outside, and reserve personnel can often observe brownish spiders descending from somewhere in the ceiling on a web. Usually they are males traveling in search of females. Their networks are often found behind furniture, in corners, under wooden panels leaning against a house – as it shall be for house spiders. But still steatoda bipunctata remains a “wild” species – it is not encountered in big cities, and if it is met a residential house, that is because its habitat is not far from it. Pachygnatha male Steatoda bipunctata female Spiders of the Lapland reserve A young Tetragnatha extensa female in autumn coloring Asagena phalerata male 19 Lapland state nature biosphere reserve Th un w an co pa Spiders of the Lapland reserve Crustullina guttata For a close genus Asagena wandering males are also very typical. They are often found on dry open slopes in the forest-tundra belt. Brightly colored spiders with thick covers are not afraid to run in a day time as they resemble big red ants very much. On open slopes of Chuna tundra in Saidnotlag tracts the tiniest species of web spiders from the local fauna are found – Crustullina guttata. Two millimeter round spiders, brown in white dots, live in the bottom surface of stones on the edges of stone falls where in the cavities between soil and stones they are constructing a rare web typical for their family consisting of a pattern raised above a substrate and many short sticky trapping threads connecting with it intended to catch small crawling insects. If you turn a stone, sun blinded crustullina will freeze on a stone surface as tiny polka dots, hoping that they will remain unnoticed, and somehow everything will return to its initial stage. Female lay only a few (5–8) huge eggs comparing to its size and its large offspring shall shed several times to grow. Therefore, the breeding cycle of crustullina is little connected with a certain season, and under one stone spiders of all ages can be found. Population of Crustullina guttata in its habitats is large; Meanwhile, in the reserve crustullina is found only in one place. This is an example of extreme confinement to a particular type of a certain biotope, and as a local settlement. It means that crustullina can not be met at the same habitat simply because they didn’t get there in sufficient amount and were not able to stay there. For the majority of spiders in the different situation is common: if there is a certain habitat certainly there are typical species inhabiting the area. The most abundant species have even “additional” habitats, where the population can be quite large, but there is a constant migration from basic habitats where population of a species lives successfully. So the picture of living environment is blurry, such as of the wolf spider Alopecosa aculeata. The most numerous Theridiidae in the reserve are inhabitants of mosses and plant litter of the genus Robertus: 7 species of small gray spiders are found in a variety of wet places. Robertus lividus under old plank Cryptachaea riparia and Theridion varians 20 ABCGheritage – Our common arctic heritage th lo on on an fin se In The most common is Robertus lividus, inhabiting somewhere under decaying trunk or old woods, their webs are coexisting with white hyphae. Several colorful species of Theridiidae inhabit also trunks and branches of trees, preferring thick fir crowns. The most common is Theridion varians, which network has no expressed pattern and consists of random loose and sticky threads. Very similar to theridion is the only one species found in the reserve from the family Mimetidae – Ero furcata. Only if you look closely, you will notice unusual rows of thin strong spines on its front legs. Before us is terrible predator – spider preying on other spiders. Ero is slowly wandering over surface of soil and plants, grabbing another spiders with its spiny limbs. If it finds a spider’s web, it starts to pull it, simulating a trapped insect; intrigued host deathwards rushes eagerly to alleged prey. In August and September somewhere in a dark corner not far from soil surface Ero female constructs a cocoon and protects it. Golden ball covered by a loose shell sits on a long thin stem, recalling a strange fruit. It contains only 6–10 large eggs. Some other species from the Linyphiidae family also have equipped front legs, however, they still hunt for insects. In late summer Drapetisca socialis abundantly inhabiting tree trunks closes its spiked limbs cabin around a prey and makes it unable to escape from a “cage”. These spiders often settle near each other, that is why Drapetisca received its species name – “social”. Such clusters can be seen, for example, at the summerhouse near Pouce cordon. Tiny Maso sundevalli and Minicia marginella hunt in the same manner in litter. Generally speaking, the Lapland reserve – is a kingdom of the Linyphiidae family, there are not less than one hundred and fifty species here – more than a half of the entire fauna. Such abundance of Linyphiidae is typical for the north Drapetisca socialis female and male Portrait of Pityohyphantes phrygianus female (up) and young Pityohyphantes male (down) Spiders of the Lapland reserve Ero furcata female and its cocoon 21 Lapland state nature biosphere reserve Helophora insignis: web and its hostess and the midland. They are second best in number of species (about 4 500) in the world after jumping spiders. Almost any spiders family can be characterized by a certain direction of evolution, a kind of unifying “idea.” Such an idea for the Linyphiidae is size reduction, miniaturization. One of the two subfamilies of Linyphiidae is called like that – spiders pygmies. Species of a medium size like, for example, Pityohyphantes phrygianus and Helophora insignis, make up a small part of the family. They build dense horizontal trapping nets with often supporting threads directed upwards and downwards; a spi- Tenuiphantes alacris: web and its owner der is walking on the underside with its abdomen up and catches a fallen insect enough right through it. With decreasing of Linyphiidae size structure of their networks becomes simpler; many spiders Pygmies can do without traps at all. Not only construction skills disappear, but also a mating behavior became easier, in contact between the sexes sexual organs structure becomes paramount – mechanism of “a lock and a key”, which prevents different species from mating. Pedipalps of male Linyphiidae often have a very complex structure. But this, apparently, was not enough, and spiders – pygmies invented an additional mechanism of “a lock and a key”. Cou- pl un no fo sp an ab lik cu er a hi is sit al th W id ne Er th ni cu Spiders pygmies Hypselistes jacksoni, Ceratinella brevis, Gonatium rubens Spiders of the Lapland reserve m ou m th w to ci bu an id in sp Spiders-Linyphiidae Neriene clathrata, Tenuiphantes alacris, Hilaira herniosa 22 ABCGheritage – Our common arctic heritage Zornella cultrigera: female and male ple of spiders-pygmies converges head-on, then male dives under a female, and she grabs his prosoma by chelicerae, but not to eat: male cephalothorax is equipped by special grooves for female “fangs”, those pits have different shapes in different species. In addition to this groups of bizarre-shaped bristles and outgrowths can develop; cephalothorax of males can look absolutely fantastic. Very often eyes area is raised up and looks like a small turret. Latin names of the species often reflect peculiarities of males’ shapes, for example, Walckenaeria monoceros. However, to describe the structure of Linyphiidae thorax a separate book would be required. So, if a male has a “key” – his pedipalps corresponding to female genitals, female “key” is her chelicerae suitable to male thorax. Tens of Linyphiidae genera living in the reserve create diversity of species comparable to variety of the rest of the families all together. Long-legged colorful tiny cob-web spiders from the genus Tenuiphante (6 species) and casting around rambling Walckenaeria (15 species); tiny whitish moss inhabitants (Palliduphantes antroniensis) and short-legged black armored Ceratinella, easily surviving drying of a loose leaf litter; beach-lovers Erigone and resident of ant hills Evansia merens; the smallest of the spiders living in the reserve – tiny Minyriolus pusillus; colonial Savignia product and Kikimora palustris; ubiquitous Zornella cultrigera; tundra Gonatium rubens and many others ... Among them there are spring, early summer, late summer and autumn species; many are encountered throughout the whole season, and zooids of several species become mature under snow; in early May, when snow starts melting they are already on the peak of the breeding season. In sunny weather, males can run directly on snow from one thaw hole to another; temperature in a few degrees above zero is sufficient for their activity. There are cases when pygmy spiders built their tiny webs right in the thick of loose spring snow and trapped migrating springtails. It is not surprisingly that being so cold resistant the Linyphiidae prevail among the northern spiders; two species even live in the zone of the Arctic polar deserts. This feature allowed species from the previous ice age to survive in the population of the reserve: population which was wide spread at that time still inhabits the tops of the Salnye tundras. Unlike the other hills of the reserve the entire range of Salnye tundras is circumcised on the southern side by a sharp almost vertical cliff starting at high altitudes, about 800–900 meters above sea level. To the north, on the contrary, elevation decreases smoothly, practically forming a plateau. The plateau is isolated by a southern cliff and from the opposite side – by the distance and poor heating of northern slopes from massive penetration of forest spiders. Salnye tundras were the only place where Collinsia holmgreni, Erigone tirolensis, Meioneta nigripes, Walckenaeria clavicornis and other species were found that are currently common in lowland tundra far to the north from the reserve. Many of them have a dark, almost black color, which is intended for more effective warm of spider’s bodies by sun. As it was mentioned above, the main body of the silk glands in the course of the evolution of spiders moved from spinnerets to an abdomen. In the most families, the foremost pair of spinnerets disappeared as unnecessary (for weaving primarily tip of abdomen, i.e. rear spinnerets are used), but some spiders have kept remaining silk glands, which plunged into an abdomen completely. Now they open outwards on special sieve-like surface called cribellum. Thread obtained from these wide sieves has a loose structure and is used for weaving distinctive lace catching nets. Earlier, all these spiders were refers to a single group of cribelliae, but recently an opinion was presented that unrelated species could keep spider gland. As a result, scientists-taxonomists started another shuffling between the families, which is still in progress – until experts come to a consensus. Representatives of the two small families in the Lapland reserve belong to spider with cribellum. Small grayish Dictyna spiders from the Dictynidae family can be met on tree trunks (Dictyna uncinata) or branched inflorescences of herbs (Dictyna major), where they build modest irregular nets with a silver delicate lace. South of the Kola peninsula in April-May population of dictyna can be huge, dried inflorescences of last year plant such as yarrow on meadows and steppes are entirely braided by them. Spiders of the Lapland reserve nd ses ll. erck die. es u- 23 Lapland state nature biosphere reserve Walckenaeria unicornis male looking for female Dictyna uncinata before hibernation Dictyna uncinata is trying to take wing Spiders of the Lapland reserve Arctobius inhabit in the lower belt of Salnye tundra Fast Mastigusa arietina having no cribellum is absolutely not similar to Dictyna. It has been recently moved into the family Dictynidae. As well as another unusual Dictynidae outstanding Hackmania prominula, Mastigusa is very rare in the reserve. The second cribellum family are Amaurobiidae represented by the only one species detected in the reserve – Arctobius agelenoides. This is quite a big and powerful spider looks like wolf spiders, but only males tend to wander. Females and baby spiders spend all their life in spinnerules between the stones. Such spinnerules came from a braid lining holes of Amaurobiidae ancestors; the mouth of the tube is disorderly framed disorderly by horizontal network, signaling about emergence of preys (a significant proportion of this family diet is constituted by beetles) – spider jumps out, grabs its prey and pulls it into the tube. Their hunting method and type of webs are ancient, many groups of primitive spiders have the same lifestyle in the tropics. Baby spiders released from a cocoon remain in the shelter for a while and after death of a female eat her body, and then they spread; life cycle takes at least two years. This large species with the most ancient way of life is found only in the lower part of the tundra nelt in the northern slope of Salnye tundras. Arctobius male 24 ABCGheritage – Our common arctic heritage
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