Spider wasp

Last updated

Spider wasp
Temporal range: Ypresian–Recent
Spiderwasp1 feb09.jpg
Western Australian pompilid captures a large huntsman spider
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Superfamily: Pompiloidea
Family: Pompilidae
Latreille, 1804
Subfamilies

see text

Wasps in the family Pompilidae are commonly called spider wasps, spider-hunting wasps, [1] or pompilid wasps. [2] The family is cosmopolitan, with some 5,000 species in six subfamilies. [3] Nearly all species are solitary (with the exception of some group-nesting Ageniellini [4] ), and most capture and paralyze prey, though members of the subfamily Ceropalinae are kleptoparasites of other pompilids, or ectoparasitoids of living spiders. [5]

Contents

In South America, species may be referred to colloquially as marabunta or marimbondo, though these names can be generally applied to any very large stinging wasps. Furthermore, in some parts of Venezuela and Colombia, it is called matacaballos, or "horse killers", while in Brazil some particular bigger and brighter species of the general marimbondo kind might be called fecha-goela/cerra-goela, or "throat locker".

Morphology

Like other strong fliers, pompilids have a thorax modified for efficient flight. The metathorax is solidly fused to the pronotum and mesothorax; moreover, the prothorax is best developed in Pompilidae and Scoliidae because wasps in these families use their forelegs to dig. [6]

A spider wasp Wasp June 2013-2a.jpg
A spider wasp

Pompilids typically have long, spiny legs; the hind femur is often long enough to reach past the tip of the abdomen. The tibiae of the rear legs usually have a conspicuous spine at their distal end. The first two segments of the abdomen are narrow, giving the body a slender look. The pompilid body is typically dark (black or blue, sometimes with metallic reflections), but many brightly colored species exist. From a lateral view, its pronotum looks rectangular and it extends back to the tegulae, near the base of the wings. Most species are macropterous (having long wings), but a few brachypterous (short-winged) and apterous (no wings) species are known. [3]

Spider wasps are best distinguished from other vespoid wasps in having (in most species) a transverse groove bisecting the mesopleuron (the mesepisternal sclerite, a region on the side of middle segment of the thorax above the point where the legs join). They have antennae with 10 flagellomeres in females and 11 in males. Most Pompilidae have straight inner eye margins. The hind wings do not have a distinct claval lobe, but they have a distinctive jugal lobe. The hind leg has a tibial spur with a tuft or row of fine hairs. The legs are long and slender with the tips of the tibia (metatibia) long enough to extend beyond the tip of the abdomen (metasoma). Sexual dimorphism is not pronounced, although females are often larger than the males. Coloration and wing appearance vary greatly among the many species. General coloration is aposematic (warning off predators), generally based on black, often with markings of orange, red, yellow, or white. [5] Larvae can also be identified by physical examination.

Systematics

The Pompilidae have in the past been split into either 4 or 6 subfamilies. However, Pitts, Wasbauer & Von Dohlen (2005) found that Notocyphinae was nested within the Pompilinae, while Epipompilinae was nested within the Ctenocerinae. This left 4 subfamilies as monophyletic clades, with Ceropalinae being the most basal subfamily and Pepsinae being the sister clade to the more derived Ctenocerinae and Pompilinae. [3] Waichert et al (2015) resurrected the Notocyphinae as a subfamily, to have 5 subfamilies within Pompilidae. In this classification Ctenocerinae were found to be the basal group; Pepsinae and Notocyphinae were sister taxa, as were Pepsinae and Pompilinae. They placed the genus Epipompilus in the Pepsinae. [7]

Notocyphus dorsalis Notocyphus dorsalis.jpg
Notocyphus dorsalis

The subfamilies according to Waichert et al are: [7]

Evolution

The oldest fossil currently known is an indeterminate fossil from the Early Eocene (Ypresian) Klondike Mountain Formation of Washington State, USA. [10] Other fossil species are known from Dominican and Baltic ambers, the Florissant Formation and various other localities in Germany, France and Spain. [11] Bryopompilus described from the mid Cretaceous Burmese amber was initially thought to belong to this family; however, it was subsequently placed in its own family, the Bryopompilidae. [12]

Ecology and behavior

Unlike many other families in the Aculeata, essentially all wasps in this family are solitary (nests made by a single female). [13]

Adult pompilids are nectar-feeding insects and feed on a variety of plants. Depending on genus and species, pompilids capture a variety of spiders for their larvae to feed on, covering nearly all free-living spider families, including tarantulas, wolf spiders (Lycosidae), huntsman spiders (Sparassidae), jumping spiders (Salticidae) and baboon spiders (Harpactirinae), though any given pompilid tends to attack only a limited diversity of spiders.

A female wasp searches the ground and/or vegetation for a spider, and upon finding one, stings it, paralyzing the spider. The targeted spider is typically unable to kill the wasp, because the wasp can just fly out of reach, so at best the spider fights fiercely to escape. [14] Tarantula hawks (Pepsini) do not attack when adult tarantulas are close to or in their burrows. Instead, the wasps seek out adult males who have left their burrows in search of females to mate with. In the open the wasp first uses its wings to beat air over the tarantula, deceiving the tarantula into thinking that it is being targeted by a large bird so the tarantula reacts by curling up to appear smaller and less noticeable, which in turn makes the tarantula defenseless against the wasp's attack. However, Brazilian Wandering spiders ( Phoneutria ) and their predators have a different interaction dynamic, and the spiders often manage to defeat the hunting wasp.[ citation needed ]

Once the spider is paralyzed, a female pompilid digs a burrow or flies or drags the spider to a previously made burrow. [15] Because of the large body size of their prey, tarantula hawks usually will either construct burrows near the site of attack or use the host's own burrow or tunnel. Pompilids typically provide each of their larvae with a single prey/host, which must be large enough to serve as its food source throughout its development. Typically, a single egg is laid on the abdomen of the spider, and the nest or burrow is closed so the larva can develop without disruption by other parasites or scavengers. [15] The female wasp may then engage in spreading soil or other changes to the area, leaving the nest site inconspicuous. One species of spider wasp protects its nests by putting dead ants into the outermost chamber, where the ants' chemicals deter predators. [16] [17]

Wasp dragging a spider to its nest Wasp and spider 02.jpg
Wasp dragging a spider to its nest

The egg hatches and the larva feeds on the spider, breaking through the integument with its mandibles. As the larva feeds on its host, it saves the vital organs, such as the heart and central nervous system, for last. By waiting until the final larval instar, it ensures the spider will not decompose before the larva has fully developed. [18] The larva has five instar stages before it pupates; no major morphological differences are noted between the first four instars, with the exception of size. At the conclusion of the final instar, the larva spins a durable silk cocoon, and emerges as an adult either later in the same season or overwinters, depending on the species and the time of year the larva pupates. [19] Some ceropalines lay their egg on a still-active spider, only temporarily paralyzing it, and the wasp larva feeds externally by extracting hemolymph after the egg hatches. In time, the spider will die, and the mature wasp larva will then pupate. [6]

The size of the host can influence whether the wasp's egg will develop as a male or a female; larger prey often yield the (larger) females. [20] Pepsis thisbe of the southwestern United States exhibits a direct correlation between adult wasp body length and the weight of its host spider, Aphonopelma echina. Because the size of a P. thisbe adult is determined by the size of the host provided for it by its mother, the seasonal frequency of host sizes implicitly will determine the size variation in adult wasps. [19]

In another study on Pepsis thisbe, [21] chemosensory cues were shown to be used to detect specific hosts. Specific chemosensory cues attract the wasp to its prey, Aphonopelma echina, despite other host spiders of the same size and frequency being present. In studies on Pepsis grossa (formerly P. formosa), [22] a pompilid of the southwestern United States, the wasps were found to have behavioral plasticity. Their hunting behavior concerning their host Rhechostica echina improved with experience. The time required to complete all behavioral components decreased with each spider killed.

Concerning mating behavior, males acquire perch territories to scan for incoming receptive females. In studies on the tarantula wasp Hemipepsis ustulata, [23] larger males are more likely to acquire perch territories and territorial males appear to increase their chances of mating because receptive females fly to perch sites held by said males.

Spider-hunting wasp Anoplius dragging a spider larger than herself backwards across a sandy heath (four stages are shown in the composite image) Pompilid Wasp Anoplius cf infuscatus dragging spider.jpg
Spider-hunting wasp Anoplius dragging a spider larger than herself backwards across a sandy heath (four stages are shown in the composite image)

Sting

Toxins

The Pompilidae produce a venom, delivered when they sting, containing a variety of powerful neurotoxins named pompilidotoxin (PMTX). These inhibit the inactivation of voltage-gated sodium channels, causing too much sodium to flow through neuron cell membranes, causing long bursts of nerve impulses (action potentials), and thus overstimulating these nerves. Alpha- and beta-PMTX are both small peptide chains of just 13 amino acids; alpha-PMTX has been studied in Anoplius samariensis, while beta-PMTX has been studied in Batozonellus maculifrons. [24] [25] [26]

Schmidt pain index

In 1984, Justin O. Schmidt developed a hymenopteran sting pain scale, now known as the Schmidt sting pain index. In this index, a 0 is given to a sting from an insect that cannot break through human skin, a 2 is given for intermediate pain, and a 4 is given for intense pain. The scale rates stings from 78 different species in 42 different genera. [27] The species Pepsis grossa, one of the species of tarantula hawk, has a sting rating of 4. [28] The sting is described as "blinding, fierce, and shockingly electric. A running hair dryer has been dropped into your bubble bath." [29]

Video of spider wasp with prey (23s)

Related Research Articles

<span class="mw-page-title-main">Velvet ant</span> Family of wasps

Velvet ants (Mutillidae) are a family of more than 7,000 species of wasps whose wingless females resemble large, hairy ants. Their common name velvet ant refers to their resemblance to an ant, and their dense pile of hair, which most often is bright scarlet or orange, but may also be black, white, silver, or gold. Their bright colors serve as aposematic signals. They are known for their extremely painful stings,, and has resulted in the common name "cow killer" or "cow ant" being applied to the species Dasymutilla occidentalis. However, mutillids are not aggressive and sting only in defense. In addition, the actual toxicity of their venom is much lower than that of honey bees or harvester ants. Unlike true ants, they are solitary, and lack complex social systems.

<span class="mw-page-title-main">Tarantula hawk</span> Common name for two genera of wasps

A tarantula hawk is a spider wasp (Pompilidae) that preys on tarantulas. Tarantula hawks belong to any of the many species in the genera Pepsis and Hemipepsis. They are one of the largest parasitoid wasps, using their sting to paralyze their prey before dragging it to a brood nest as living food; a single egg is laid on the prey, hatching to a larva which eats the still-living host. They are found on all continents other than Europe and Antarctica.

<span class="mw-page-title-main">Wasp</span> Group of insects

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. The wasps do not constitute a clade, a complete natural group with a single ancestor, as bees and ants are deeply nested within the wasps, having evolved from wasp ancestors. Wasps that are members of the clade Aculeata can sting their prey.

<span class="mw-page-title-main">Pompilinae</span> Subfamily of wasps

The Pompilinae are a subfamily of the spider wasp family, Pompilidae, the species of which lay their eggs on the paralyzed bodies of their prey.

<span class="mw-page-title-main">Pepsinae</span> Subfamily of wasps

The Pepsinae are a subfamily of the spider wasp family, Pompilidae, including the two genera of large tarantula hawks, as well as many genera of smaller species.

<i>Anoplius nigerrimus</i> Species of wasp

Anoplius nigerrimus is a species of spider wasp, or pompilid, and is the type species of the genus Anoplius.

<i>Epipompilus insularis</i> Species of wasp

Epipompilus insularis is a species of spider wasp which is endemic to New Zealand and it is the only species of the genus Epipompilus found in New Zealand.

Evagetes crassicornis is a kleptoparasitic spider wasp with a holarctic distribution.

<i>Hemipepsis ustulata</i> Species of wasp

Hemipepsis ustulata is a species of tarantula hawk wasp native to the Southwestern United States. Tarantula hawks are a large, conspicuous family of long-legged wasps that prey on tarantulas by using their long legs to grapple with their prey and then paralyze them with a powerful sting. They are solitary, displaying lekking territorial behavior in their mating rituals.

<i>Anoplius viaticus</i> Species of wasp

Anoplius viaticus, commonly known as the black-banded spider wasp, is a species of spider wasp. These wasps are known as spider wasps because the females capture spiders to provide their offspring with food. The paralysed spider is cached in a burrow, the wasp lays an egg on it, and when this hatches, the developing wasp larva consumes the spider. This species is found in sandy heathland across most of Europe.

Anoplius infuscatus is a species of spider wasp found mainly in Eurasia.

Ireangelus is a genus of kleptoparasitic spider wasps from the sub-family Ceropalinae of the family Pompilidae. The genus has a pan tropical distribution, being known from Oriental, Neotropical, Australian, eastern Palearctic, and Madagascan Zoogeographic regions being best represented in the Neotropics. Irenangelus is closely related to the more widespread genus Ceropales, the two genera forming a monophyletic subfamily, Ceropalinae within the Pompilidae. This is regarded as the most basal grouping of the Pompilidae but this view is problematic because of the kleptoparasitic life history of the Ceropalines, it is now considered that they Ceropalines and other pompilids evolved from a common ectoparasitoid ancestor.

<i>Tachypompilus ferrugineus</i> Species of wasp

Tachypompilus ferrugineus, the rusty spider wasp, red-tailed spider hunter, or sometimes red-tailed spider wasp is a species of spider wasp from the Americas. It preys mainly on wandering spiders, especially wolf spiders.

<i>Poecilopompilus algidus</i> Species of wasp

Poecilopompilus algidus is a species of spider wasp which is widespread in the Americas.

<i>Pepsis grossa</i> Species of wasp

Pepsis grossa is a very large species of pepsine spider wasp from the southern part of North America, south to northern South America. It preys on tarantula spiders, giving rise to the name tarantula hawk for the wasps in the genus Pepsis and the related Hemipepsis. Only the females hunt, so only they are capable of delivering a sting, which is considered the second most painful of any insect sting; scoring 4.0 on the Schmidt sting pain index compared to the bullet ant's 4.0+. It is the state insect of New Mexico. The colour morphs are the xanthic orange-winged form and the melanic black winged form. In northern South America, a third form, known as "lygamorphic", has a dark base to the wings which have dark amber median patches and a pale tip.

<i>Allochares azureus</i> Species of wasp

Allochares azureus is a species of spider wasp from the family Pompilidae, it is the only member of the monotypic genus Allochares. It occurs in the southern part of North America and is a specialist parasitoid of the Southern house spider.

<i>Notocyphus</i> Genus of spider wasps

Notocyphus is a genus of spider wasps, belonging to the family Pompilidae. They are the only genus in the monotypic subfamily Notocyphinae. These wasps are found in the Nearctic and the Neotropics.

<i>Ceropales bipunctata</i> Species of wasp

Ceropales bipunctata, the Two-Speckled Cuckoo Spider Wasp, is a species of diurnal, kleptoparasitic spider wasp in the family Pompilidae. It is an obligate kleptoparasite, meaning that it must rely on the captured provisions of other spider wasps and cannot capture its own. It is found on the Atlantic Coast of North America, north to New Brunswick, Canada. It feeds on nectar primarily from goldenrods, and also other common nectaring plants. It is known to lay an egg on the prey of other pompilids, including the two species Anoplius cleora and Anoplius aethiops. Eggs are laid in the book lung of the spider. These wasps also sometimes lay an egg on prey from spider-hunting wasps in the family Sphecidae.

<span class="mw-page-title-main">Ageniellini</span> Tribe of insects

Ageniellini, known as the mud-nesting spider wasps, is a tribe of spider wasps in the subfamily Pepsinae.

<i>Pepsis mildei</i> Western Hemisphere tarantula-hawk wasp

Pepsis mildei, also known as Milde's tarantula-hawk wasp, is a species of predatory spider wasp native to the Western Hemisphere. These wasps capture live tarantulas to feed to their larva; the adults graze on flowers. Tarantula hawks generally have no meaningful predators. The wing color is black and/or orange, with rare leucistic wing coloration known in males. The wings of P. mildei are duller overall than those of Pepsis grossa or Pepsis cinnabarina. Per Edward Essig, Milde's tarantula-hawk wasp usually measures 20–30 millimeters in length and is a metallic blue-black overall, with "the antennae reddish, dusky at extreme base in the male and the basal third dusky in the female; the wings fiery red with the bases and apices dusky". In all tarantula wasps, the antennae can be used to distinguish between males and female: males are straight or gently arced, while the females have curved antennae. The lifespan of males is one to two months, while female Milde's tarantula-hawk wasps live for a longer span of time.

References

  1. ↑ "Pompilidae Spider-Hunting Wasp". NBN Atlas. Retrieved 10 August 2019.
  2. ↑ Costa, Fernando G.; Pérez-Miles, Fernando; Mignone, Antonio (April 2004). "Pompilid Wasp Interactions with Burrowing Tarantulas: Pepsis cupripennis versus Eupalaestrus weijenberghi and Acanthoscurria suina (Araneae, Theraphosidae)" (PDF). Studies on Neotropical Fauna and Environment. 39 (1): 37–43. doi:10.1080/01650520412331270945. ISSN   0165-0521. OCLC   231044349. S2CID   84475545 . Retrieved 20 April 2012.
  3. 1 2 3 Pitts, James P.; Wasbauer MS & von Dohlen CD (2005). "Preliminary morphological analysis of relationships between the spider wasp subfamilies (Hymenoptera: Pompilidae): revisiting an old problem". Zoologica Scripta. 35 (1): 63–84. doi:10.1111/j.1463-6409.2005.00217.x. S2CID   86180326.
  4. ↑ Evans, H.E.; Shimizu, A. (1996). "The evolution of nest building and communal nesting in Ageniellini (Insecta: Hymenoptera: Pompilidae)". Journal of Natural History. 30 (11): 1633–1648. doi:10.1080/00222939600770961.
  5. 1 2 Goulet, H; Huber, JT, eds. (1993). Hymenoptera of the world: an identification guide to families. Agriculture Canada. pp. 202–205.
  6. 1 2 Daly, Howell V.; Doyen, John T.; Purcell, Alexander H. (1998). Introduction to Insect Biology and Diversity. Oxford: Oxford UP. ISBN   978-0-19-510033-4.
  7. 1 2 Waichert, Cecilia; Rodriguez, Juanita; Wasbauer, Marius; von Dohlen, Carol & Pitts, James (2015). "Molecular phylogeny and systematics of spider wasps (Hymenoptera: Pompilidae): Redefining subfamily boundaries and the origin of the family". Zoological Journal of the Linnean Society. 175 (2): 271–287. doi: 10.1111/zoj.12272 .
  8. 1 2 3 Loktionov, Valery M. (2023). "Geographical distribution of the spider wasps (Hymenoptera: Pompilidae) of the world". Russian Entomological Journal. 32 (4): 394–402. doi: 10.15298/rusentj.32.4.05 .
  9. ↑ "Classification and checklist of Afrotropical Pompilidae". Archived from the original on 2012-01-27. Retrieved 2012-03-18.
  10. ↑ Archibald, S. B.; Rasnitsyn, Alexandr P.; Brothers, Denis J.; Mathewes, Rolf W. (2018-01-08). "Modernisation of the Hymenoptera: ants, bees, wasps, and sawflies of the early Eocene Okanagan Highlands of western North America". The Canadian Entomologist. 150 (2): 205–257. doi:10.4039/tce.2017.59. ISSN   0008-347X. S2CID   90017208.
  11. ↑ Rodriguez, Juanita; Waichert, Cecilia; von Dohlen, Carol D.; Pitts, James P. (2017-10-11). Wong, William Oki (ed.). "The geological record and phylogeny of spider wasps (Hymenoptera: Pompilidae): A revision of fossil species and their phylogenetic placement". PLOS ONE. 12 (10): e0185379. Bibcode:2017PLoSO..1285379R. doi: 10.1371/journal.pone.0185379 . ISSN   1932-6203. PMC   5636076 . PMID   29020022.
  12. ↑ Rodriguez, Juanita; Waichert, Cecilia; von Dohlen, Carol; Poinar Jr., George; Pitts, James (2015). "Eocene and not Cretaceous origin of spider wasps (Pompilidae): fossil evidence from amber". Acta Palaeontologica Polonica. doi: 10.4202/app.00073.2014 .
  13. ↑ CSIRO Fact Sheet
  14. ↑ "Death Match: Tarantula vs. Wasp". 10 April 2014.
  15. 1 2 Spider Wasps Australian Museum Online
  16. ↑ "Dead-ant wall protects young spider wasps". Science News. 2014-07-07. Retrieved 2014-07-11.
  17. ↑ Staab, Michael; Ohl, Michael; Zhu, Chao-Dong; Klein, Alexandra-Maria (2014-07-02). "A Unique Nest-Protection Strategy in a New Species of Spider Wasp" (PDF). PLOS ONE. 9 (7): e101592. Bibcode:2014PLoSO...9j1592S. doi: 10.1371/journal.pone.0101592 . PMC   4079592 . PMID   24987876.
  18. ↑ Punzo, F (2005). "Studies on the natural history, ecology, and behavior of Pepsis cerberus and P. mexicana (Hymenoptera: Pompilidae) from Big Bend National Park, Texas". Journal of the New York Entomological Society. 113 (1): 84–95. doi:10.1664/0028-7199(2005)113[0084:sotnhe]2.0.co;2. S2CID   84879231.
  19. 1 2 Punzo, F (1994). "The biology of the spider wasp, Pepsis thisbe (Hymenoptera: Pompilidae) from Trans Pecos, Texas. I. Adult morphometrics, larval development and the ontogeny of larval feeding patterns". Psyche: A Journal of Entomology. 101 (3–4): 229–242. doi: 10.1155/1994/70378 .
  20. ↑ "Spider predators and parasites". Iziko Museums of Cape Town: museums.org.za. Retrieved 28 November 2006.
  21. ↑ Punzo, F.; Ludwig, L. (2005). "Behavioral Responses of Pepsis thisbe (Hymenoptera: Pompilidae) to Chemosensory Cues associated with host spiders". Journal of Insect Behavior. 18 (6): 757–766. doi:10.1007/s10905-005-8738-0. S2CID   22280009.
  22. ↑ Punzo, F.; Garman, B. (1989). "Effects of encounter experience on the hunting behavior of the spider wasp, Pepsis formosa (Say) (Hymenoptera: Pompilidae)". The Southwestern Naturalist. 34 (4): 513–518. doi:10.2307/3671510. JSTOR   3671510.
  23. ↑ Alcock, J.; Kemp, J. (2006). "The behavioral significance of male body size in the Tarantula Hawk Wasp Hemipepsis ustulata (Hymenoptera: Pompilidae)". Ethology. 112 (7): 691–698. doi:10.1111/j.1439-0310.2006.01204.x.
  24. ↑ Kawai N, Konno K (2004). "Molecular determinants of two neurotoxins that regulate sodium current inactivation in rat hippocampal neurons". Neurosci. Lett. 361 (1–3): 44–6. doi:10.1016/j.neulet.2004.01.006. PMID   15135889. S2CID   38819369.
  25. ↑ Sahara Y, Gotoh M, Konno K, Miwa A, Tsubokawa H, Robinson HP, Kawai N (2000). "A new class of neurotoxin from wasp venom slows inactivation of sodium current". Eur J Neurosci. 12 (6): 1961–70. doi:10.1046/j.1460-9568.2000.00084.x. PMID   10886337. S2CID   34486331.
  26. ↑ Schiavon, Emanuele; Stevens, Marijke; Zaharenko, André J.; Konno, Katsuhiro; Tytgat, Jan; Wanke, Enzo (2010). "Voltage-gated sodium channel isoform-specific effects of pompilidotoxins". FEBS Journal. 277 (4): 918–930. doi:10.1111/j.1742-4658.2009.07533.x. ISSN   1742-464X. PMID   20059541.
  27. ↑ Berenbaum, M (2003). "A Stinging Commentary". American Entomologist. 49 (2): 68–69. doi: 10.1093/ae/49.2.68 .
  28. ↑ Schmidt, Justin O. (1990). "Hymenoptera Venoms: Striving Toward the Ultimate Defense Against Vertebrates". In D. L. Evans; J. O. Schmidt (eds.). Insect Defenses: Adaptive Mechanisms and Strategies of Prey and Predators. Albany, New York: State University of New York Press. pp. 387–419. ISBN   0-88706-896-0.
  29. ↑ Schmidt, Justin (2016). The Sting of the Wild. Baltimore, Maryland: Johns Hopkins University Press. ISBN   978-1-4214-1929-9.
This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.