Tigrosa Helluo
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''Tigrosa helluo'' is a species of spider belonging to the family
Lycosidae Wolf spiders are members of the family Lycosidae (). They are robust and agile hunters with excellent eyesight. They live mostly in solitude, hunt alone, and do not spin webs. Some are opportunistic hunters, pouncing upon prey as they find it or ...
, also known as wolf spiders. ''T. helluo'' was formerly known as ''Hogna helluo'' before differences between dorsal color patterns, habitat preferences, body structures, etc. were discovered. The species is native to the United States, Canada, and Mexico. It can be found across the eastern half of the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
, primarily in the Northeast and
New England New England is a region comprising six states in the Northeastern United States: Connecticut, Maine, Massachusetts, New Hampshire, Rhode Island, and Vermont. It is bordered by the state of New York to the west and by the Canadian provinces ...
, and as far west as
Nebraska Nebraska () is a state in the Midwestern region of the United States. It is bordered by South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Kansas to the south; Colorado to the southwe ...
and
Kansas Kansas () is a state in the Midwestern United States. Its capital is Topeka, and its largest city is Wichita. Kansas is a landlocked state bordered by Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to the ...
. ''T. helluo'' can be found in diverse habitats including woods, marshes, fields, and riparian areas. Typically, members of this species prefer to live in wetter areas as opposed to dry environments. Males tend to live for around a year and females will live for close to two years. The defining characteristic of ''T. helluo'' is its brown
carapace A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...
and distinct yellow stripe starting from its anterior eyes extending down the
cephalothorax The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. (The terms ''prosoma'' and ''opisthosoma'' are equivalent to ''cepha ...
. The underside of the
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
has distinct black spots that distinguish ''T. helluo'' from other wolf spiders. At an average size of around , ''T. helluo'' is one of the smaller wolf spiders. In addition, females tend to be larger than males in a display of
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
. Like other spiders of the family Lycosidae, wolf spiders are solitary and live and hunt alone. Unless in the process of mating, ''T. helluo'' will remain alone. It is usually active at night, during which it hunts for prey. In addition, ''T. helluo'' does not create webs, although females will make burrows under rocks or boards. Relying on their excellent eyesight and senses, ''T. helluo'' members are great hunters. Members of ''T. helluo'' are not very aggressive and will not attack humans unless provoked. Their bites inject venom; however, it is not medically significant to humans, as only minor pain and swelling occur.
Antivenom Antivenom, also known as antivenin, venom antiserum, and antivenom immunoglobulin, is a specific treatment for envenomation. It is composed of antibodies and used to treat certain venomous bites and stings. Antivenoms are recommended only if th ...
is rarely needed.


Description

Whereas average body length for '' Tigrosa'' can range from for females and for males, ''T. helluo'' body length averages . Compared to its close relative '' T. aspersa'', ''T. helluo'' is much smaller. ''T. helluo'' is also often mistaken for ''
Pisaurina mira ''Pisaurina mira'', also known as the American nursery web spiders, is a species of spider in the family Pisauridae. They are often mistaken for wolf spiders (Lycosidae) due to their physical resemblance. ''P. mira'' is distinguished by its uniqu ...
'', the nursery web spider, due to the physical resemblance. However, the carapace of ''T. helluo'' consists of a brownish color contrasting with a distinctive yellow stripe that extends from the anterior eyes to the cephalothorax. Starting from the posterior median eyes, a set of fainter yellow stripes extends posteriorly. In addition, the underside of the abdomen is marked with several black spots. Patterns found on the dorsal side of the cephalothorax and abdomen are similar to those found on ''T. georgicola''; however, the faint yellow stripes appearing on ''T. helluo'' do not extend as far. Black spots found on the abdomen of ''T. helluo'' are also not found on ''T. georgicola''. Coloration in the legs differs between sexes, with males' legs appearing yellow without distinct markings and females' legs appearing reddish brown without additional markings or bands.


Taxonomy

The ''Tigrosa'' genus includes four species—'' T. annexa'', '' T. aspersa'', '' T. grandis'', and ''T. helluo''—that were moved from the genus ''
Hogna ''Hogna'' is a genus of wolf spiders with more than 200 described species. It is found on all continents except Antarctica. Etymology The word ''Hogna'' might be a rough latinization of one of the Greek words '' ὄχνη'' (''ókhnē'') "p ...
''. ''Tigrosa'' is defined by the distinct colored pattern on the dorsal cephalothorax, structure of the male palpus and epigynum, leg length, foraging behaviors, and eye arrangement. The aggressive nature and colorful patterns of these spiders related to the meaning of the word ''Tigrosa'', which is "fierce like a tiger".Allen R. Brady "Nearctic species of the new genus Tigrosa (Araneae: Lycosidae)," The Journal of Arachnology 40(2), 182-208, (1 August 2012). https://doi.org/10.1636/K11-77.1


Habitat and distribution


Habitat

In the United States, ''T. helluo'' can be found in a large variety of states in woods,
marsh A marsh is a wetland that is dominated by herbaceous rather than woody plant species.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p Marshes can often be found at ...
es, and even grassy areas in northeastern states such as
Connecticut Connecticut () is the southernmost state in the New England region of the Northeastern United States. It is bordered by Rhode Island to the east, Massachusetts to the north, New York to the west, and Long Island Sound to the south. Its cap ...
. Females' nests are often found under stones or boards in fields and woods. These nests consist of burrows lined with
silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the coc ...
. However, in states like Florida and
Mississippi Mississippi () is a state in the Southeastern region of the United States, bordered to the north by Tennessee; to the east by Alabama; to the south by the Gulf of Mexico; to the southwest by Louisiana; and to the northwest by Arkansas. Miss ...
, ''T. helluo'' is most often found in wetter areas such as marshes and
lake A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much large ...
s.


Distribution

''T. helluo'' can be found throughout the United States, Canada, and Mexico. It primarily resides along the East Coast in the New England region but can be found along the entire East Coast down to
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
. ''T. helluo'' resides in most states in the eastern half of the United States. It can be found as far west as Kansas and Nebraska.


Reproduction and life cycle

Mating for ''T. helluo'' usually occurs in May or June. Males will live for just over one year while females will often live for around two years.Balfour, R. (2004). Differences in the growth of the wolf spider Hogna helluo (Araneae: Lycosidae) reared under high and low food quantity diets. (Electronic Thesis or Dissertation). Retrieved from https://etd.ohiolink.edu/ Maturation occurs in males between May and September. Egg sacs are made in May and July and can range from in diameter. The eggs by themselves can appear tan or yellow and are around in diameter. Females will carry these eggs in an egg sac on their back until they hatch. Even after hatching, the female will continue to carry the hatchlings on her back until they mature enough to hunt by themselves. Female maturation can occur at any point during the year. Once mature, females will begin to
overwinter Overwintering is the process by which some organisms pass through or wait out the winter season, or pass through that period of the year when "winter" conditions (cold or sub-zero temperatures, ice, snow, limited food supplies) make normal activi ...
during colder months. Males will typically die before it gets too cold. As ''T. helluo'' matures, it will
molt In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
periodically. Time and duration of molting is dependent on nutritional intake and consequent growth potential. The frequency of molting is based on the nutritional reserves of an individual ''T. helluo''. Individuals that feed more often will molt less frequently during maturation.


Mating


Female–male dimorphism

''T. helluo'' is
sexually dimorphic Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
. In this species, the two sexes dramatically differ in size, with mature females being larger than males. Typically, females will grow to weigh more than with carapace widths of around , while males will usually only weigh in excess of and have carapace widths of around .


Sexual cannibalism

Like many other wolf spiders, ''T. helluo'' will engage in precopulatory and postcopulatory
sexual cannibalism Sexual cannibalism is when an animal, usually the female, cannibalizes its mate prior to, during, or after copulation.Polis, G.A. & Farley, R.D. Behavior and Ecology of Mating in the journal of Arachnology 33-46 (1979). It is a trait observed in m ...
. The frequency at which females participate in sexual cannibalism depends on a variety of factors. The degree of sexual dimorphism between two spiders has been shown to have an effect on the occurrence of sexual cannibalism. Males who are smaller are more vulnerable to larger females and therefore are more likely to be cannibalized. The larger the size difference between the male and female, the more likely for sexual cannibalism to occur. Thus, sexual cannibalism in ''T. helluo'' can be viewed similarly to a predator–prey interaction where size difference affects the actions of each. Another factor that impacts the degree of sexual cannibalism for ''T. helluo'' is the female condition. Females who have been starved and therefore are hungry are more likely to cannibalize than females who are well fed. It is possible that males can maximize their fitness by mating only with well-fed females. The male condition can also affect the possibility of sexual cannibalism. Older males or males in poorer condition, while they might not be very small, are more vulnerable to females. Therefore, they are more likely to alter their copulatory behavior depending on their perceived risk of being cannibalized. These males can invest more into a single copulation by engaging for longer because they are at a high risk of being eaten after mating.


Copulation

The process of
copulation Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetra ...
for ''T. helluo'' consists of a series of interactions between the male
palp Pedipalps (commonly shortened to palps or palpi) are the second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") ...
and the female epigynum. First, the male will mount the female such that they are facing opposite from each other. The underside of the male's
cephalothorax The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. (The terms ''prosoma'' and ''opisthosoma'' are equivalent to ''cepha ...
is positioned against the dorsal section of the female's abdomen. The male then will signal the female to rotate the abdomen by touching the anterior of the female abdomen. After this, the male will use his palp to engage with the female's abdomen and epigynum. The right and left palps will interact with the right and left sides of the epigynum. After the palp and epigynum are engaged, the male will expand his hematodocha, causing his
embolus An embolus (; plural emboli; from the Greek ἔμβολος "wedge", "plug") is an unattached mass that travels through the bloodstream and is capable of creating blockages. When an embolus occludes a blood vessel, it is called an embolism or emb ...
to enter the female copulatory duct.
Semen Semen, also known as seminal fluid, is an organic bodily fluid created to contain spermatozoa. It is secreted by the gonads (sexual glands) and other sexual organs of male or hermaphroditic animals and can fertilize the female ovum. Semen i ...
is then exchanged. This interaction is called an ''insertion''. During a single insertion, the male may also expand his hematodocha multiple times. However, it is often only a single expansion per insertion. ''T. helluo'' will also engage in more than one insertion per copulation. Usually, one insertion will happen per side of the male palp. Copulation for ''T. helluo'' will typically last around eight minutes.


Diet


Prey

Wolf spiders typically eat different kinds of
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s or even other smaller spiders like '' Pardosa milvina''. They are nocturnal so they wait for prey and hunt at night. ''T. helluo'' can survive on insects such as
crickets Crickets are orthopteran insects which are related to bush crickets, and, more distantly, to grasshoppers. In older literature, such as Imms,Imms AD, rev. Richards OW & Davies RG (1970) ''A General Textbook of Entomology'' 9th Ed. Methuen 8 ...
, fly grubs,
cockroaches Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known a ...
,
mealworm Mealworms are the larval form of the yellow mealworm beetle, ''Tenebrio molitor'', a species of darkling beetle. Like all holometabolic insects, they go through four life stages: egg, larva, pupa, and adult. Larvae typically measure about or ...
s, and even
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s.


Effects of diet on survivorship

''T. helluo'' is able to feed on a large variety of prey and thus enjoys a mixed diet of different insects. A 1992 study assessing the effects of polytypic versus monotypic diets on wolf spiders found that ''T. helluo'' individuals raised on polytypic diets had significantly higher survivorship and reached sexual maturity earlier than individuals consuming a monotypic diet. Certain body parts like the cephalothorax and legs were also significantly larger at maturity in the polytypic diet group.


Effects of diet on behavior

A 1999 study examining the effect of feeding on burrow construction for ''T. helluo'' reported that hunger level significantly impacts its behavioral decisions. In comparison to starved spiders, well-fed individuals had a better nutritional state and made significantly more burrows. Since males do not make burrows, they were not included in the study.


Predators

Although they are hunters, wolf spiders are also preyed upon by other organisms such as
praying mantises Mantises are an order (Mantodea) of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae ("mantids"). Mantises are distributed worldwide in temperate and tropical habitats. They ha ...
, birds,
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s, small reptiles, and even other
spider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s. Lycosidae have fantastic vision and are sensitive to vibrations, so they use these senses to detect and evade predators. They also use their speed and camouflage from their natural coloring to avoid predators. Females of the species ''T. helluo'' make burrows in secluded habitats, such as underneath stones, that can also function as a place to hide from predators. If needed, ''T. helluo'' can also bite predators and inject
venom Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a sti ...
.


Behavior


Hunting

''T. helluo'' typically switches between a few different foraging sites. Females build burrows of silk and wait for prey to show up in their current foraging site. Males typically roam around and wait for prey. Like all wolf spiders, ''T. helluo'' is a solitary hunter and will hunt alone, meeting with other spiders only to mate.


Chemical cues in hunting

A 2001 study examining chemical cues in ''T. helluo'' suggests that ''T. helluo'' is able to detect chemical cues from recently hunted prey. The spiders will show a preference for chemical cues that match that of their most recently hunted prey. Thus, further hunting activities will show a dietary preference for the prey species most recently consumed. It has also been shown that the presence of
glyphosate Glyphosate (IUPAC name: ''N''-(phosphonomethyl)glycine) is a broad-spectrum Herbicide, systemic herbicide and Crop desiccation, crop desiccant. It is an organophosphorus compound, specifically a phosphonate, which acts by inhibiting the plan ...
-based herbicide affects prey capture behavior of ''T. helluo''. In the presence of herbicide, ''T. helluo'' is better able to detect and orient itself towards prey and therefore is able to subdue prey more efficiently.


Communication

While wolf spiders are solitary and do not usually associate with others outside mating, communication between individual spiders does occur occasionally. Wolf spiders have some of the best vision of all spiders, so they use visual cues, such as waving their pedipalps, to signal to each other for mating. They are also sensitive to vibrations, scent, and taste.


Bites


Bites to humans

Wolf spiders are usually not aggressive and will not bite unless readily provoked. They also possess venom that is injected into organisms upon biting. However, to humans, wolf spider bites are generally considered minor and not medically significant. Bites typically cause minor swelling and pain, but no severe complications.
Antivenom Antivenom, also known as antivenin, venom antiserum, and antivenom immunoglobulin, is a specific treatment for envenomation. It is composed of antibodies and used to treat certain venomous bites and stings. Antivenoms are recommended only if th ...
is not usually needed to treat wolf spider bites.


Differences from ''Pardosa milvina''

Both ''Pardosa milvina'' and ''Tigrosa helluo'' are among the more common wolf spider species in the United States, but they do have behavioral differences in their hunting approaches. Whereas ''T. helluo'' tends to wait for prey to pass by its location prior to attacking, ''P. milvina'' goes out to find its prey. However, the much smaller size of ''P. milvina'' also makes it prey for ''T. helluo''.


References


External links

* {{taxonbar, from1=Q27502863, from2=Q2299904 Lycosidae Spiders described in 1837 Spiders of North America