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Copula

In linguistics, a copula is a verb or verbal construction that links the subject of a sentence or clause to a subject complement, such as a noun, adjective, or prepositional phrase, thereby expressing relations of identity, attribution, location, or state.^[1] The term originates from the Latin word copula, meaning "link" or "bond," reflecting its role in connecting elements of a proposition.^[1] In English, the most prototypical copula is the verb to be (e.g., "The sky is blue"), which functions to equate the subject with its complement or describe its state.^[2] Other common copular verbs include seem, appear, become, feel, look, sound, smell, and taste, which often convey sensory impressions or changes in state (e.g., "She became a teacher" or "The soup tastes delicious").^[1] These verbs are intransitive, typically followed by complements rather than objects, and differ from auxiliary verbs by serving as the main verb in the clause.^[3] Copulas vary significantly across languages, with some employing multiple forms for different semantic nuances, while others omit them entirely in certain contexts. In Romance languages like Spanish, Portuguese, and Italian, copulas often derive from Latin esse ("to be"), but Ibero-Romance varieties (e.g., Spanish ser and estar) distinguish between permanent states (ser) and temporary ones (estar), a distinction influenced by Vulgar Latin verbs like stare ("to stand") and sedēre ("to sit").^[4] Many languages, including Russian, Arabic, Hebrew, and various Austronesian and Turkic languages, frequently lack an overt copula in present-tense declarative sentences, relying instead on juxtaposition of subject and predicate (e.g., Russian: "On student" meaning "He is a student").^[5] Zero copulas are also prevalent in creole languages and varieties like African American Vernacular English, where absence rates vary by predicate type (e.g., higher before adjectives and locatives than before nouns).^[6] Historically, the concept of the copula traces back to ancient philosophy and grammar, where Aristotle identified it as the connective element in judgments linking subject and predicate.^[7] In Indo-European languages, copular roots often stem from Proto-Indo-European forms such as h₁es- (related to existence or being) and steh₂- (related to standing), which evolved into modern verbs like English be, German sein, and Sanskrit asti.^[8] Missionary grammars of non-Indo-European languages, such as Aymara in the 17th century, further adapted the term to describe nominal sentences without verbs, highlighting its cross-linguistic applicability.^[9] Beyond linguistics, "copula" also denotes a logical connector in propositions (e.g., the implied "is" in "All humans are mortal"), an anatomical structure in biology connecting two parts (such as in the hyoid bone or tongue development), a genus of cnidarians, and, in statistics, a multivariate probability distribution function that captures dependence between variables while preserving uniform marginals (e.g., Gaussian or Clayton copulas). In arts and entertainment, it refers to elements like the rhythm game beatmania IIDX 23 copula.^[10]^[11] These usages underscore the term's broad application in linking disparate elements across disciplines.

Linguistics

Linking Verb

In linguistics, a copula is a verb or verbal construction that serves as a linking element between the subject and a subject complement, such as a predicate nominative or predicate adjective, without conveying an action or event. For instance, in the English sentence "The sky is blue," the verb "is" functions as the copula, equating the subject "the sky" with the adjectival complement "blue." The primary grammatical function of a copula is to identify, equate, or ascribe a property to the subject, thereby establishing a relation of predication rather than agency or process. Common copular verbs in English include "be" in its various inflections (e.g., "am," "was," "been"), as well as "seem," "become," and sensory verbs like "feel," "appear," "look," "sound," "smell," and "taste," which some analyses classify as copular when they link to a complement without implying physical action. For example, in "She seems happy," "seems" links the subject "she" to the adjective "happy," expressing a state of appearance; similarly, "The soup tastes delicious" uses "tastes" to attribute a sensory quality to the subject. These verbs typically require a complement to complete the clause and cannot stand alone as full predicates in the same way action verbs can. The term "copula" derives from Latin copula, meaning "a bond" or "that which binds together," reflecting its role in connecting grammatical elements. It entered linguistic usage in the 1640s to describe verbs that unite subjects and predicates. Cross-linguistically, copular functions often trace back to Proto-Indo-European roots like *h₁es- ("to be"), which evolved into present-tense forms such as English "is," Latin est, and Sanskrit asti. A basic example is the English "She is a teacher," where "is" equates the subject to a nominal complement; in contrast, some languages imply this linkage without an overt verb. Copulas manifest in various types across languages, including the zero copula, where no explicit verb appears and the linkage is implied by juxtaposition; for example, in Russian, "Ona uchitel'nitsa" means "She (is a) teacher," omitting the copula in present-tense predicative clauses.^[12] Similarly, Standard Arabic employs a zero copula in present-tense constructions, as in "al-bayt kabīr" ("The house (is) big"), relying on subject-complement agreement for tense and case marking.^[13] The support copula acts in an auxiliary-like role to provide morphological or syntactic support, such as English "be" in questions or negations (e.g., "Is she a teacher?" via do-support analogy). Finally, the specific copula refers to dedicated inflectional forms tailored to particular semantic or syntactic contexts, as seen in Spanish with "ser" for inherent properties (e.g., "Ella es profesora" – "She is a teacher") versus "estar" for temporary states (e.g., "Ella está cansada" – "She is tired").

Copula Omission and Variation

Copula omission, also known as zero copula, is a widespread feature in the world's languages, where no overt linking element appears between the subject and predicate in verbless clauses. Similarly, Arabic vernaculars frequently employ null copulas in present tense predicational constructions, such as Lebanese Arabic l-bornayṭa ∅ meškle ("The projector [is a] problem"), relying on word order and context to convey the linkage. This omission is not random but conditioned by factors like tense (often restricted to present) and predicate type (nominal or adjectival). According to typological surveys, approximately 45% of sampled languages permit zero copula for predicate nominals, highlighting its prevalence across families.^[12] Copula variation manifests in diverse structural types, reflecting grammatical strategies to link subjects and predicates. Inflectional copulas, which change form to mark tense, person, and number, are exemplified by the English verb "be," with forms like am, is, was, and were adapting to syntactic contexts. Periphrastic copulas, involving multi-word constructions, appear in languages like Irish, where is ea serves as a copula in equative clauses, such as Is ea an t-úll ("It is the apple"), combining a particle with a pronoun for emphasis or specificity. Additionally, some languages feature copula-like elements where adjectives function predicatively without a separate linker; in Japanese, i-adjectives like takai ("high") inflect directly as predicates, as in Sora ga takai ("The sky [is] high"), behaving verb-like without an overt copula. The historical development of copulas traces back to Proto-Indo-European roots, notably h₁es-, which yielded forms like English "is," Latin esse, and Irish is, serving as the core for existential and equative meanings across Indo-European languages. In Romance languages, this evolved into complex systems; Spanish and Portuguese distinguish ser (from Latin esse, for inherent or permanent properties, e.g., Ella es profesora "She is a teacher") from estar (from Latin stare "to stand," for temporary states, e.g., Ella está cansada "She is tired"), a split emerging around the 13th century to encode aspectual nuances. Grammaticalization processes often drive such changes, with lexical verbs (e.g., posture verbs like "sit" or "stand") bleaching into copular functions, as seen in Arabic vernaculars where forms like Libyan gāʕəd ("sitting") grammaticalize into an overt copula for contingent states, gāʕəd ɣādi ("He [is] over there"). Typological patterns reveal further diversity, including in non-spoken modalities like sign languages. In American Sign Language (ASL), copular linkage often relies on non-manual markers such as head tilts or eye gaze rather than an overt manual sign, though the sign SELF has emerged as a copula for equative and class membership predications, as in contexts denoting identity (e.g., "I [am] a linguist").^[14] Grammaticalization here follows a cycle from deictic gestures to demonstratives and then copulas. Cross-family examples abound: Niger-Congo languages like Swahili exhibit zero copula in some nominal predications, such as "Yeye mwalimu" ("He [is a] teacher"); Austronesian languages such as Tagalog similarly omit it in certain contexts, "Siya ay guro" but often without 'ay' in informal speech. These patterns underscore how copulas—or their absence—adapt to language-specific grammars, with zero copula more common in isolating or agglutinative systems.

Mathematics and Statistics

Copula Functions

In probability theory and statistics, a copula is defined as a multivariate cumulative distribution function (CDF) C: [0,1]^d \to [0,1] for which each marginal distribution is uniform on the interval [0,1]. This means that for any \mathbf{u} = (u_1, \dots, u_d) with u_i \in [0,1], C(\mathbf{u}) represents the joint probability P(U_1 \leq u_1, \dots, U_d \leq u_d), where each U_i follows a standard uniform distribution. Copulas serve as a mathematical tool to capture the dependence structure between random variables independently of their marginal distributions.^[15] The foundational result linking copulas to general multivariate distributions is Sklar's theorem, established in 1959. It states that any d-dimensional CDF F with continuous marginal CDFs F_1, \dots, F_d can be expressed as F(x_1, \dots, x_d) = C(F_1(x_1), \dots, F_d(x_d)) for some copula C, and conversely, for any copula C and marginal CDFs F_1, \dots, F_d, the function F(x_1, \dots, x_d) = C(F_1(x_1), \dots, F_d(x_d)) defines a joint CDF with those marginals. For continuous marginals, the copula is unique; in the discrete case, it is unique on the support of the marginals. This theorem underscores the role of copulas in separating the marginal behaviors from the overall dependence.^[16]^[15] Copulas possess several key properties that ensure they function as valid joint CDFs. They are grounded, meaning C(u_1, \dots, u_{j-1}, 0, u_{j+1}, \dots, u_d) = 0 for any j and \mathbf{u} \in [0,1]^d. Their marginals are uniform on [0,1], so C(u_1, \dots, u_j, 1, \dots, 1) = u_j for each j. Additionally, copulas are 2-increasing, which implies that the volume of the d-dimensional "rectangle" [0, \mathbf{u}] is non-decreasing in each argument, ensuring non-negative probabilities. Boundary conditions further specify that C(0, \dots, 0) = 0 and C(1, \dots, 1) = 1. These properties guarantee that copulas adhere to the axioms of multivariate CDFs while focusing solely on dependence.^[15] The class of all possible copulas is bounded by the Fréchet-Hoeffding bounds, which provide the theoretical limits on dependence. The upper bound, known as the comonotonic copula, is M(\mathbf{u}) = \min(u_1, \dots, u_d), representing perfect positive dependence. The lower bound, the countermonotonic copula, is W(\mathbf{u}) = \max\left(\sum_{i=1}^d u_i - (d-1), 0\right), capturing the strongest possible negative dependence (achievable in the bivariate case but not always in higher dimensions). All copulas satisfy W(\mathbf{u}) \leq C(\mathbf{u}) \leq M(\mathbf{u}) for \mathbf{u} \in [0,1]^d. These bounds, originally derived in the contexts of correlation theory, delineate the extremal dependence structures admissible under given marginals.^[15] Common examples illustrate these concepts. The independence copula is the product \Pi(\mathbf{u}) = \prod_{i=1}^d u_i, which models no dependence between variables. The comonotonic copula M exemplifies complete positive association. The Gaussian copula arises from the multivariate normal distribution with mean zero and correlation matrix \Sigma, defined as C_{\Sigma}(\mathbf{u}) = \Phi_{\Sigma}(\Phi^{-1}(u_1), \dots, \Phi^{-1}(u_d)), where \Phi is the standard univariate normal CDF and \Phi_{\Sigma} is the multivariate normal CDF. This copula is widely used due to its flexibility in capturing linear correlations via \Sigma.^[15]

Dependence Modeling

Copulas provide a flexible framework for modeling dependence in multivariate data by linking univariate marginal distributions to their joint distribution, enabling the isolation and analysis of complex dependence structures that go beyond linear measures like the Pearson correlation coefficient, which assumes linearity and fails to capture non-linear relationships or tail dependence in extreme events.^[17] This separation allows practitioners to model scenarios where variables exhibit asymmetric or extreme co-movements, such as joint defaults in financial portfolios or concurrent floods in hydrological systems.^[17] Several families of copulas are employed to represent diverse dependence patterns. Archimedean copulas, constructed using a generator function \phi(t), include the Clayton copula, C(u,v;\theta) = \left( u^{-\theta} + v^{-\theta} - 1 \right)^{-1/\theta} for \theta > 0, which emphasizes lower tail dependence suitable for modeling shared downside risks, and the Gumbel copula, C(u,v;\theta) = \exp\left( -\left[ (-\ln u)^{\theta} + (-\ln v)^{\theta} \right]^{1/\theta} \right) for \theta \geq 1, which captures upper tail dependence for extreme co-occurrences.^[17] Elliptical copulas, derived from multivariate elliptical distributions, encompass the Gaussian copula for symmetry and the t-copula, which incorporates heavy tails through degrees of freedom parameter \nu, making it ideal for datasets with clustered extremes as detailed in Demarta and McNeil (2005).^[18] For high-dimensional settings, vine copulas decompose the multivariate dependence into a cascade of bivariate copulas, facilitating flexible modeling of intricate structures, as introduced by Aas et al. (2009). Dependence measures derived from copulas include rank correlations and tail coefficients. Kendall's tau is computed as \tau = 1 - 4 \iint [C(u,v) - uv] \, du \, dv, providing a robust, distribution-free assessment of monotonic dependence.^[17] Spearman's rho, given by \rho = 12 \iint C(u,v) \, du \, dv - 3, quantifies concordance based on ranks and aligns with percentile-based associations.^[17] Tail dependence coefficients, such as the upper tail \lambda_U = \lim_{u \to 1^-} P(U_2 > u \mid U_1 > u) = 2 - \lim_{u \to 1^-} \bar{C}(u,u)/(1-u), where \bar{C} is the survival copula, measure the conditional probability of joint extremes, essential for risk assessment in tails.^[17] Estimation of copula parameters typically involves maximum likelihood methods for parametric families, assuming known or estimated marginals, or non-parametric approaches like the empirical copula via the Rosenblatt transform, which maps data to uniform pseudo-observations.^[19] Goodness-of-fit tests, such as the Cramér-von Mises statistic based on the empirical process of transformed ranks, evaluate model adequacy by comparing observed and fitted dependence, with asymptotic distributions derived for inference.^[20] The application of copulas in dependence modeling gained prominence following Sklar's 1959 development, particularly in finance where they underpin collateralized debt obligation (CDO) pricing via Gaussian or t-copulas to simulate default correlations, as pioneered by Li (2000); post-2008 financial crisis analyses highlighted their role in underestimating tail risks during market crashes.^[21] In hydrology, copulas model joint flood risks by capturing spatial and temporal dependencies between river discharges, enabling multivariate frequency analysis for dam design and risk mitigation, as demonstrated by Favre et al. (2004).^[22]

Biology

Anatomical Structures

In anatomy, a copula refers to a narrow ligamentous or bony connection between two structures, serving as a linking element in various bodily systems.^[23] For instance, the body of the hyoid bone functions as a copula, uniting its greater and lesser horns to form a stable U-shaped framework in the anterior neck.^[23] A prominent example in embryology is the copula linguae, a mesenchymal swelling that arises in the floor of the pharynx during the fourth week of gestation from the mesoderm of the second pharyngeal arch.^[24] This structure, also known as the copula or part of the hypobranchial eminence, contributes to the formation of the posterior third of the tongue by overgrowing the earlier tuberculum impar and integrating contributions from the third and fourth pharyngeal arches between weeks 5 and 7.^[25] The copula linguae fuses with the hypobranchial eminence, a median swelling inferior to it derived from the third and fourth arches, ensuring the posterior tongue's development while the anterior two-thirds forms separately from the first arch's lateral lingual swellings.^[26] This fusion process establishes the tongue's dual innervation, with the posterior region supplied by the glossopharyngeal nerve (cranial nerve IX) from the third arch.^[25] In the skeletal system, copular connections appear in the hyoid apparatus, where lateral ligaments such as the thyrohyoid ligaments link the hyoid bone's greater horns to the superior horns of the thyroid cartilage, facilitating laryngeal elevation during swallowing.^[27] In comparative vertebrate anatomy, similar structures are evident; for example, in fish, the copula forms part of the hyoid arch as a ventral cartilaginous element, such as the basibranchial copula, connecting gill arches and supporting respiratory mechanics.^[28] These features are conserved across vertebrates, highlighting the copula's role in pharyngeal and branchial arch evolution.^[29] The term "copula" originates from Latin copula, meaning "a bond" or "connection," reflecting its use to describe structural joining rather than reproductive copulation.^[30] In anatomical literature, it gained prominence in 19th-century embryology texts to denote such linking formations, building on earlier descriptive traditions in comparative morphology.^[31] Disruptions in copula development can lead to congenital anomalies; for instance, incomplete degeneration of tissues during tongue separation from the pharyngeal floor may result in ankyloglossia (tongue-tie), characterized by a short lingual frenulum that restricts tongue mobility and affects feeding or speech.^[25] This condition arises from persistent fusion remnants in early embryogenesis, including elements influenced by the copula's integration.^[32]

Cnidarian Genus

Copula is a genus of small box jellyfish in the family Tripedaliidae, class Cubozoa, within the phylum Cnidaria. The genus was established in 2010 to accommodate species previously classified under Carybdea that possess distinctive adhesive pads on the exumbrella for attachment to substrates.^[33] Originally with the sole species Copula sivickisi, the genus was expanded in 2024 to include Copula lucentia sp. nov., described from the western Mediterranean Sea.^[34] Copula sivickisi, originally described as Carybdea sivickisi by Stiasny in 1926 and reclassified into Copula based on phylogenetic and morphological analyses, remains the type species.^[33] This reclassification helps maintain the monophyly of related genera and highlights evolutionary relationships among cubozoans.^[35] New distribution records for C. sivickisi include first observations in Philippine waters as of 2025.^[36] Physically, C. sivickisi is a diminutive cubomedusa with a bell height of up to 12 mm (width up to 14 mm) and a transparent, cube-shaped body typical of box jellyfish.^[37] It features four slender tentacles equipped with nematocysts—specialized stinging cells used for capturing prey—and four unique adhesive pads at the bell's apex that enable it to cling to surfaces like seaweed or rocks during rest. The species exhibits sexual dimorphism, with males and females distinguishable by subtle differences in gonad structure and tentacle morphology.^[38] Copula sivickisi inhabits coastal waters of the Indo-Pacific region, including areas around Japan, Australia, and Hawaii, though records extend to the tropical western Atlantic and Indian Oceans.^[39] Unlike larger, pelagic box jellyfish such as those in the genus Chironex, it prefers nearshore, reef-associated environments and is often found in mangroves or harbors at depths of less than 5 meters.^[40] Its distribution reflects a metapopulation structure influenced by ocean currents and behavioral adaptations that limit dispersal.^[39] Behaviorally, C. sivickisi is nocturnal, hiding via adhesive pads during the day and actively foraging at night on small crustaceans like copepods and zoea larvae.^[41] It employs vision to navigate toward bioluminescent signals from prey-rich areas, extending its tentacles to ensnare victims.^[42] The species' venom, delivered through nematocysts, is far less potent than that of Irukandji syndrome-causing jellyfish, typically resulting only in mild skin irritation or blistering in humans rather than severe envenomation.^[43] Reproduction occurs in the medusa stage through internal fertilization following a unique courtship ritual where males and females entangle tentacles to transfer sperm, a behavior that inspired the genus name "Copula," meaning "link" in Latin. First documented in the 1990s through surveys in Hawaii and Japan, C. sivickisi serves as a model organism in studies of cubozoan evolution, particularly regarding vision, mating systems, and phylogenetic systematics.^[40]^[33]

Arts and Entertainment

Music

In medieval music theory, particularly within the polyphonic practices of the 13th century, a copula denotes a specific contrapuntal texture characterized by a lower voice—typically the tenor—holding long, sustained notes derived from plainchant, while the upper voices introduce faster-moving, ornamental lines in measured rhythm. This style creates a layered contrast between the stable foundation of the tenor and the more florid duplum or triplum above it, serving as a transitional form between freer organum and stricter discant. The term originates from the Latin copula, meaning "coupling" or "bond," which aptly describes the interconnected yet differentiated voicing that "links" the parts in early polyphony.^[44]^[45] The copula emerged prominently in the motets and organa of the Notre Dame school in Paris during the late 12th and early 13th centuries, where it facilitated rhythmic experimentation within liturgical frameworks. Composers such as Pérotin employed copula sections in works like his four-voice organum Viderunt omnes (c. 1198), integrating them to enhance expressiveness during melismatic passages on vowels, allowing upper voices to vocalize rapidly against the tenor's holds. This technique is evident in the Magnus Liber Organi, the foundational collection attributed to the school, and in conductus pieces where copula provides ornamental relief. Manuscripts like the Montpellier Codex (c. 1270–1300) preserve numerous motets featuring copula textures, showcasing its role in bridging chant-based polyphony with emerging secular influences.^[44]^[46] Theoretical analysis of the copula appears in key treatises of the era, notably Johannes de Garlandia's De mensurabili musica (c. 1260), which defines it as an intermediate style "between organum and discant," emphasizing its rhythmic mode application to upper parts while the tenor remains unmeasured or broadly sustained. This distinction highlights the copula's departure from pure organum's melismatic freedom and discant's equal-voiced equality, focusing instead on textural hierarchy. By the 14th century, copula elements evolved into more intricate isorhythmic motets, as seen in the works of Philippe de Vitry, where sustained tenors combined with repeating rhythmic patterns in upper voices, influencing the Ars Nova's structural innovations. Unlike rhythmic "copulatio"—a term for specific ligature groupings in notation—the copula prioritizes overall voice coupling over isolated rhythmic units.^[47]^[45]^[44]

Video Games

Beatmania IIDX 23: Copula is an arcade rhythm video game developed and published by Konami, released exclusively in Japan on November 11, 2015, as the 23rd installment in the long-running Beatmania IIDX series that originated in 1999.^[48] The title draws its name from the Latin word for "link" or "coupling," thematically evoking the connection of musical beats and train cars through its artwork and slogan, "Every day shines," which ties into a train journey motif across seasonal and mystery unlock lines.^[48] It features 91 new original songs, bringing the total library to 1,002 tracks, including 17 special LEGGENDARIA charts designed for advanced players with enhanced difficulty levels up to 12.^[49] Building on the core BEMANI franchise mechanics, Copula retains the seven-key gameplay system where players press buttons and a turntable in time with on-screen arrows to match rhythm tracks across single or double-play modes.^[50] Key innovations include the introduction of HELL CHARGE NOTES, which function as dual-value holds that rapidly deplete the life gauge if missed, adding tension to high-speed sequences; the ASSISTED EASY modifier, which lowers the clear threshold from 80% to 60% for accessibility; and an expanded ALL BEGINNER folder with numerical difficulty ratings for easier entry-level play.^[48] The game also revives BPM and difficulty-based song folders, introduces EX and MIRROR options for dan courses (graded challenge modes), and upgrades double-play dan courses to four songs starting at 7-kyu rank, with STEP UP mode now scaling up to level 9 based on prior performance.^[51] Hardware compatibility remains with standard IIDX arcade cabinets, supporting e-AMUSEMENT online features for score tracking and unlocks until their discontinuation on November 30, 2016.^[48] Culturally, Copula reinforced the IIDX series' prominence in Japanese arcades, particularly among rhythm game enthusiasts, with its train-themed unlocks fostering replayability through seasonal events like Season Line and Mystery Line, where players collect virtual stamps to access hidden tracks.^[48] The game's soundtrack, featuring diverse genres from electronic to hard dance, was released internationally on platforms like iTunes, extending its reach to global fans who created custom charts and playthrough videos despite the arcade-only format.^[52] While specific installation numbers are not publicly detailed, its integration of no new licensed music—marking a shift toward originals—highlighted Konami's focus on in-house production amid the series' enduring legacy in the rhythm genre, influencing subsequent titles without direct sequels to this entry.^[48] Beyond Copula, the term appears sparingly in video game contexts, such as indie mods or mechanics emphasizing "linking" elements in puzzle or rhythm simulations, though no other major titles prominently feature it in name or core theme.^[53]

References

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[32]
Evolution of box jellyfish (Cnidaria: Cubozoa), a group of highly toxic ...
Nov 18, 2009 · The new genus Copula is defined to contain tripedaliids that possess adhesive pads on the exumbrellar apex with which they attach themselves to ...
[33]
Evolution of box jellyfish (Cnidaria: Cubozoa), a group of highly toxic ...
Nov 18, 2009 · In Copula sivickisi, a mature male and female engage in sexual activity by entangling their tentacles. While swimming as a couple, the male ...
[34]
[PDF] Box Jellyfish (Cubozoa: Carybdeida) in Hawaiian Waters, and the ...
Dec 4, 2014 · rEMarKS: Copula sivickisi is a small sexually dimorphic cubomedusa ≤14 mm bell diameter with four distinctive adhesive pads that allow ...Missing: 1999 | Show results with:1999
[35]
(PDF) Gonadal histology of box jellyfish (Cnidaria: Cubozoa) reveals ...
Mar 31, 2018 · This comparative histological study examines the gonad organization of Alatina alata and Copula sivickisi, two box jellyfish species that ...
[36]
Behavioural and oceanographic isolation of an island-based ...
May 13, 2021 · The Copula sivickisi metapopulation covers tropical and temperate latitudes in the Pacific, Indian and Atlantic Oceans. While this expansive ...
[37]
Copula - Jellipedia
Copula catches zooplankton at night, while during the day it uses adhesive pads to attach to seaweed. Males and females of the species can be identified based ...
[38]
Vision in the Nocturnal Box Jellyfish Copula sivickisi - PMC - NIH
Mar 30, 2016 · The animals use vision when hunting. When they are active at night they seek out high prey-concentration by visual attraction to areas with abundant ...Missing: taxonomy | Show results with:taxonomy
[39]
[PDF] jiggling - NIWA
Fortunately, the sting of Copula sivickisi is not known to be fatal to humans, although it can leave a blister- like wound. Copula sivickisi is the only box ...
[40]
Notre Dame (Chapter 27) - The Cambridge History of Medieval Music
In a sense it combines the attributes of organum and conductus: the motet is based on a bit of plainchant in the tenor, but the voices above it carry newly ...
[41]
The "Copula" According to Johannes de Garlandia - jstor
Reckow's musical example and discussion of this sentence (Die Copula, pp. 17 ? 19) suggest that he has misunderstood this. Significantly, his reprinting of.
[42]
The Montpellier Codex, Part 4 - A-R Editions
1-day delivery 30-day returnsThe musical transcriptions are included in parts I–III; part IV provides text commentary and translations, an index of first lines, and guides to pronunciation.Missing: copula | Show results with:copula
[43]
Johannes de Garlandia Concerning Measurable Music (c.1260) | PDF
Johannes de Garlandia Concerning Measurable Music (c.1260). This document discusses different types of measurable music, including discant, copula, and organum.
[44]
beatmania IIDX 23 copula - RemyWiki
Release Information · Announcement date: July 8th, 2015 · Location test dates: July 10th to 12th, 2015 · Release date: November 11th, 2015 · Slogan: 毎日が、輝く ...
[45]
beatmania IIDX 23 copula full song list - RemyWiki
2.21 beatmania IIDX 2nd style (8 songs); 2.22 beatmania IIDX substream (8 songs); 2.23 beatmania IIDX (6 songs). 3 LEGGENDARIA (17 songs). New Songs (91 songs).
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beatmania IIDX 23: copula (2015) - MobyGames
Apr 19, 2020 · The gameplay is still the same as in beatmania, with the addition of two extra keys.Missing: features | Show results with:features
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History of Notable Changes in beatmania IIDX - IIDX.org
beatmania IIDX 23 copula (November 11, 2015). Added HELL CHARGE NOTE; Added ASSIST EASY gauge; Added ALL BEGINNER folder; DP class mode has 4 songs – previously ...
[48]
Beatmania IIDX 23 Copula (Original Soundtrack) - Apple Music
Feb 24, 2016 · Disc 2 ; 1. Donkey Donk · Ryu ; 2. Wheel of Journey · Vivian ; 3. Serotonin · U1 overground ; 4. Alice in Mystic Garden · DJ TOTTO ; 5. Taiyo Sun Sun ...
[49]
Beatmania IIDX 23 (Copula) location testing | NeoGAF
Jul 11, 2015 · New modifier introduced: ASSIST-EASY, which lowers clear border from 80% to 60%. Using this counts as an ASSIST CLEAR if a song is cleared.