The Cambridge Dictionary of Human Biology and Evolution Larry L

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Abdur Reef

A. n dates: see Oakley’s dating series in box below. A antigen: epitope that specifies the A in the ABO AAA: abbreviation for several societies of interest to blood group. It consists of four precursor sugars human evolutionary biologists, including the attached to glycoproteins of the cell membrane, American Anthropological Association and the aka H substance, plus a specific fifth terminal sugar, American Anatomical Association. N-acetylgalactosamine, that is attached by an enzyme. A Oakley’s absolute dating series A.1 date: highest of Oakley’s hierarchical levels of absolute dating, the direct dating of a specimen, e.g. by measuring the radiocarbon activity of a bone itself. A.2 date: one of Oakley’s hierarchical levels of absolute dating, dating derived from direct determination by physical measurement of the age of the sediments containing the fossil specimen. A.3 date: one of Oakley’s hierarchical levels of absolute dating, the correlation of a fossil-bearing horizon with another deposit whose age has been determined directly by A.1 or A.2 methods. See biostratigraphy. A.4 date: lowest of Oakley’s hierarchical levels of absolute dating, estimating an absolute age on the basis of some theoretical consideration, such as matching climatic fluctuations observed in strata with astro- nomically derived curves of effective solar radiation, or matching terrestrial glacial and interglacial episodes with the known marine paleotemperature or oxygen isotope stage.

(Cf. R.n dates)

This enzyme differs from its homologue that attaches abdominal circumference: anthropometric meas- the B-specific sugar by one amino acid substitution. urement; distance around the torso measured with See B antigen and O null allele. a tape measure placed at the level of the greatest AAFS: see American Academy of Forensic anterior extension of the abdomen in the horizon- Scientists. tal plane, normally the umbilicus, passed horizon- AAPA: see American Association of Physical tally around the body. The measurement is taken Anthropologists. after a normal expiration. Used for various body abaxial: on the opposite side of, or facing away from, indices pertaining to adipose distribution. Cf. waist the axis of a structure or an organism. circumference. Abbeville: archaeological site found in 1830 in the abdominal skinfold: anthropometric measurement; French Valley of the Somme, possibly dated to the skinfold measured about 3 cm lateral to the umbil- first interglacial (ϭ Pleistocene deposits) described icus. Used in combination with other skinfold by Boucher de Perthes in 1847; one of the first his- measurements to estimate body composition. See torical cases of artifacts of arguable antiquity; used skinfold thickness. as proof of the existence of early humans contrary to abductor: muscle or muscle group that moves a body the story contained in Genesis. Through the early part away from the mid-line; aka levator. Cf. adductor. twentieth century the term Abbevillian was applied abductor pollicis brevis: intrinsic muscle of the globally to the most ancient tool deposits; Oakley hand; it originates from the scaphoid and trapezium lumped both Oldowan choppers and Acheulean and inserts into the lateral surface of metacarpal I. Its choppers under this term. Aka Port du Bois, action is to abduct the thumb; one of the thenar Menchecourt-les-Abbeville. group of muscles. Cf. adductor pollicis. Abbevillian tool tradition: term that refers to pre- abductor pollicis longus: muscle that acts on the Acheulean stone-tool assemblages found in Europe; hand; it originates from the posterior surface of the among the artifacts are crude bifacial tools con- radius and ulna and inserts into the base of structed with a hammerstone rather than a softer metacarpal I. Its action is to abduct and extend the striking instrument (as in Acheulean).Aka Chellean. thumb. See Oldowan. Abdur Reef: archaeological site near Abdur along the abdomen: portion of the body between the thorax and Bari Coast in Eritrea on the coast of the Red Sea, the pelvis. dating to 125kya (U/Th). The elevated reef contains abdominal cavity: space within the body between the artifacts including bifacial hand-axes, blade tools respiratory diaphragm and the pelvic inlet in which and sharpened obsidian flakes found among the abdominal viscera are contained; abdomen. shellfish remains and animals bones deposited on an Cf. coelom. uplifted reef system. Aka Abdur Limestone Site.

© Cambridge University Press www.cambridge.org Cambridge University Press 0521664861 - The Cambridge Dictionary of Human Biology and Evolution Larry L. Mai, Marcus Young Owl and M. Patricia Kersting Excerpt More information

abris

aberrant: 1. a departure from the norm; an anomaly or antecedent to later colonizers. In reference to humans, aberration. 2. in genetics, an individual phenotype this term has been applied most often to the original that exhibits atypical characteristics due to the inhabitants of Australia; aborigine. Cf. indigenous, influence of the environment rather than from autochthonous. genetic factors. abortion: 1. arrest of a process or disease before com- ABH blood group (ABH): a polymorphism found in pletion. 2. interruption of pregnancy before the fetus many organisms and determined by the presence or has attained a stage of viability, usually before the absence of A, B, and/or H antigens found on cell 24th gestational week. There are two classes: an membranes such as erythrocytes. In humans, this induced abortion is the termination of a pregnancy, system has been modified and is named the ABO often for medical reasons, by means other than those blood group. causing spontaneous abortion (aka clinical abortion, abiotrophy: 1. general term denoting degenerative therapeutic abortion, feticide). A spontaneous abortion changes of tissue due to genetic causes. 2. deterior- is the natural termination of a pregnancy before the ation of the body. fetus can survive outside the uterus (between 0 and abnormal: 1. not normal. 2. departing from the usual 20 weeks); occurs in about 30% of first pregnancies, structure, position, or condition. Abnormality. frequently so early that a woman is not aware that abnormal hemoglobins: any of the hemoglobin she was pregnant, and is most often due to chromo- alleles other than hemoglobin A; most are causes of somal errors (50–60%); sometimes applied to a hemoglobinopathies, such as sickle cell anemia specific period of gestation, such as during the first and thalassemia. 20 weeks of pregnancy (aka early fetal death). See ‡ ABO blood group: polymorphism based on the miscarriage and stillbirth. presence or absence of two antigens (IA and IB) found Abri Pataud: archaeological site found in 1949 near on the cell membrane of erythrocytes (red blood Les Eyzies in the Vézère River Valley, Dordogne, cells); the antigens are also found on other cells. The France, dating from 34kya and persisting for over ‘O’ factor is an absent or ‘null’ allele, not an antigen. 14 ky; excavated by H. Movius; contains over 25000 See H substance. Aurignacian II artifacts including burins, an engrav- ABO disease association: any abnormal condition ing tool for working antlers, a venus figurine in relief correlated with an allele or genotype of the ABO on stone, and hominid remains from at least two blood group. A number of diseases are associated individuals, including a young female attributed to with the presence of each of the A, B and O alleles in Homo sapiens. Of 30 crates of animal bones excav- the ABO blood group; among the more prominent ated, about 30% of all prey appear to have been rein- are salivary gland tumors (type A), ulcers (type O), deer. Aka Pataud. and certain infant diarrheas (types A and B). Abri Suard: archaeological site found in 1870 in the La ABO hemolytic disease of the newborn (HDN): Chaise cave area, western France, dated to 200–100 destruction of the erythrocytes of a fetus; a mal- kya; contains hominid remains from several indi- adaptive condition roughly twice as common as the viduals including an incomplete 1050cm3 calotte Rhesus incompatibility form of HDN, but almost assigned to the archaic Neanderthal group. Aka always clinically milder in its manifestation. Caused Suard. by some of the smaller anti-A or anti-B antibodies in Abrigo do Lagar Velho rockshelter: archaeological a mother’s immune system penetrating the placen- site found in 1998 in the Lapedo Valley, north of tal membranes. Lisbon, Portugal, dated to 24.5kya (radiocarbon), ABO incompatibility: lowered probability of concep- that contains Gravettian artifacts. The fossil tion owing to the genotypes in the ABO blood group hominid remains include a nearly complete skeleton possessed by potential parents. In populations that of a child (Lagar Velho 1) attributed to either Homo maintain all three alleles, it has been suggested sapiens or H. neanderthalensis. Among the arti- that type O mothers produce fewer offspring than facts collected were charcoal, stone tools, pierced deer expected when the fathers are either type A or type B; canines and fossilized red deer bones and horse this may be due to the presence of either antibodies teeth. Aka Lagar Velho. This find has been inter- in the mother’s vaginal secretions (that could agglu- preted by some as indicative of gene flow between tinate sperm), or certain smaller classes of maternal Neandertals and AMHs, and in support of the anti-A and anti-B antibodies that could pass through multiregional continuity model. See pierced the placenta and affect the development of the fetus. mammal teeth and last of the Neandertals.Cf. aboriginal: indigenous; native to; original biological Mezmaiskaya Cave. species or populations inhabiting a particular geo- abris: French for shelter; rockshelter is ‘abris sous graphical region, particularly referring to groups roche’.

acculturation

Absarokius Matthew, 1915: poorly known tarsiiform more accurate than counting radioactive decay (con- belonging to the subfamily Anaptomorphinae, family ventional radiocarbon dating); AMS allows dating Omomyidae. Found in early to middle Eocene based upon very small samples (such as a speck of deposits from the Rocky Mountain region of North wood), and also extends the useful range of the tech- America. Three to seven species, two of which may be in nique back to almost 100kya. an ancestor – descendant relationship; Tetonius acceptor stem: that region of a tRNA molecule to homunculus is suggested as an ancestor for at least one which a specific amino acid is attached in an enzy- species. Dental formula: 2.1.3.3/2.1.3.3 or 2.1.3.3/2.1.2.3, matic reaction, which loads the tRNA prior to the depending on the species. Absarokius differs from other genetic translation step of polypeptide synthesis. anaptomorphines in the enlargement of the third and accessory olfaction system (AOS): special sensory fourth premolars and in thickening of the mandible, cells in the vomeronasal organ of the nose plus their suggesting a diet of hard morsels. Body mass estimates connections to the brain. The AOS is distinct from range from 130 to 500 g. Synonym for Anaptomorphus. the sense of smell, which is part of the main olfactory See Appendix 1 for taxonomy. system. The AOS receives social and sexual signals in abscess: localized collection of pus in a cavity due to the form of pheromones from other members of the the accumulation of white blood cells; often due to same species. infections caused by Staphylococcus bacteria in skin and ‡ acclimation: short-term plasticity of individual other tissues. phenotype; somatic physiological response by indi- ‡ absolute dating (technique): any of the methods viduals to environmental pressures. See developmen- that provide an estimate of a specific date or age in tal acclimation. Not to be confused with evolution or solar years, subject to probabilistic limits, from genetic adaptation.Cf.acclimatization. material recovered from an archaeological or pale- ‡ acclimatization: phenotypic plasticity in an ontological site. Common absolute techniques organism that enables it to make physiologic include dendrochronology, radiocarbon dating changes that reduce the strain caused by stresses (see carbon 14), and potassium – argon dating. from environmental factors. Acclimatization may Aka chronometric dating. See dating techniques. be short-term (see acclimation) or long-term (see absolute geological time: scale established by geo- developmental plasticity). It has been suggested physicists that is defined by time in years ago, thou- that the ability to respond physiologically to the sands of years ago (kya), and millions of years ago environment is partly genetic but requires environ- (mya); the time scale is derived by using absolute mental interaction to be expressed. Verb: acclima- dating techniques that depend upon the regular tize. decay of unstable elements into stable daughter accommodation: 1. responses by an organism to products. The absolute geological time scale of the environmental stress that are not wholly successful earth currently extends back over 4.5 billion years because, even though they favor survival of the indi- (by) of earth history. Aka the geological time scale. vidual, they also result in significant losses in some See Appendix 4. important functions. 2. alteration in the convexity of absolute risk: probability that an organism will the lens of the eye to increase visual acuity. experience a certain event, such as a disease; risk is accretion model: scheme in which fossil hominids calculated on the basis of test results and/or the formerly termed archaic or transitional form are fur- occurrence of the event in relatives. ther segregated into Stage 1, the early pre- absorption: process by which a substance passively Neandertals (e.g. Mauer and Petralona); Stage 2, the enters a body membrane. pre-Neandertals (e.g. Steinheim, Swanscombe, and acatalasia: genetic disorder marked by deficiency of Atapuerca); Saale – Eem Stage 3, the early the enzyme catalase; manifestations range from mild Neandertals (e.g. Biache); and Weichsel Stage 4, the (ulcers in tooth sockets) to severe (recession of tooth ‘classic’ Neandertals (e.g. La Chapelle and La sockets with gangrene of the gums). Two principle Ferrassie). This model is assumed by some to repre- forms of acatalasia have been identified: (a) Japanese sent a temporal series of a single isolated gene pool. type, and (b) Swiss type. Aka acalalesemia. accretionary growth: multiplication of components acaudate: without a tail, as in the great apes; acaudal. in an organ or tissue without qualitative functional accelerating differentiation: after isolation of popu- change. lations due to differences accrued at a few loci, the acculturation: nearly symmetrical increase in the sim- rapid increase of differentiation at many loci result- ilarity of two autonomous cultural systems, and the ing in broad geographic radiation. lessening of culture distance between them; one of accelerator mass spectrometry (AMS): method of the possible outcomes set in motion by the meeting counting single radiocarbon atoms that is much of the two systems. Cf. assimilation.

acromegaly

acculturation model: proposal advanced to explain causing rhizomelic shortening, but may also cause the 10 ky temporal overlap of Neandertals and Cro- trident hand, frontal bossing and mid-face hypopla- Magnons in central and southern Europe in which sia; the cranial base may distend, causing the cranium transitional tool industries such as the to become enlarged. The defective gene is a fibroblast Châtelperronian result from the interaction growth factor receptor. Cf. hGH-resistant dwarfism between Neandertals and anatomically modern and pituitary dwarfism. ‘invaders’ bearing an Aurignacian tool tradition. achromatopsia: see color blindness. According to this model, the development and use of acidic protein: any of certain classes of protein that decorated bone objects and body ornaments by coat a DNA molecule, binding to specific regions, and Neandertals was the result of borrowed or mimicked involved in genetic regulation. culture rather than of independent invention. acidity: 1. the quality or state of being acid measured as ‡ accuracy: 1. freedom from error. 2. degree of con- the concentration of free, unbound hydrogen ions in formity of a measured quantity to the true value of a solution. The higher the Hϩ concentration, the what is being measured, where the true value is rep- greater the acidity and the lower the pH; an acid has resented by a standard. a pH less than 7.0. 2. the acid content of a fluid. acentric chromosome: chromosome fragment that acquire: to develop after birth. Adjective: acquired. Cf. lacks a centromere. congenital, hereditary. acentric primate group: terrestrial primates that flee ‡ acquired characteristic: refers to the outdated into trees when confronted with danger. Cf. cen- Lamarckian concept of transformism, in which phe- tripedal primate group. notypic modifications arise solely through environ- acetabulocristal buttress: thickening of the hip bone mental influences on the developmental processes of between the acetabulum and the iliac crest. Found in an organism; some proponents believe that such char- bipedal hominids but not in pongids.One of the neces- acters can be passed on to the next generation, now sary modifications that led to habitual bipedalism. largely discredited. Aka use inheritance, use – disuse, acetabulum: cup-shaped socket formed from the and soft inheritance. See adaptation, Baldwin effect, junction of the ilium, ischium, and pubis on the evolution, Lamarckism, and Lysenkoism. coxal bone. The acetabulum is on the lateral surface Acquired Immune Deficiency Syndrome: see and receives the head of the femur with which it AIDS. forms a ball-and-socket joint. In female humans acquired immunity: immunity to a specific pathogen this structure is often oriented anteriorly, whereas in acquired after birth owing to exposure by either nat- males it is more lateral. Adjective: acetabular. ural or artificial means. Immunity results from the acetylcholine: ubiquitous neurotransmitter that is activation of specific B cells or T cells exposed to the involved in the transmission of signals at nerve pathogen’s antigens. This exposure results in the synapses. production of antibodies and memory cells. Aka Aché: foragers or hunter-gatherers in modern-day acquired defense. Paraguay who have survived with a subsistence ‡ acrocentric chromosome: chromosome in which lifestyle in small-scale societies into the twenty-first the centromere is very near one end so that the short century. arm is very small or minute and the long arm is much ‡ Acheulean tool tradition: stone-tool technology longer. characterized by large, pointed, almond-shaped acrocephaly: condition of being high-headed, such bifacial tools called hand-axes and cleavers, whose that the height of the skull is at least 98% of the exact purpose is unknown. Smaller flake tools were breadth. also present, but in lesser numbers. Acheulean tools acrocranic index: in reference to the cranial breadth – range in time from 1.5 mya to 200kya; widespread in height index, with an index of 98.00 or greater; such the Old World. The Acheulean industry gets its name an individual is considered to have a high skull. from St. Acheul, a site in France. In African sites, acrodont: having rootless teeth attached to the alveolar where Acheulean tools are often found at sites with ridge of the jaws, e.g. human deciduous dentition. Developed Oldowan tools at the same level, the acromegaly: form of gigantism often confused with hand-axes are usually found near sources of water several similar disorders of which acromegaly is a such as stream channels. feature; it is genetically heterogenous. Familial Achilles tendon: see calcaneal tendon. acromegaly is characterized by continued growth achondroplasia (ACH): a congenital, autosomal after a normal adolescent growth spurt, resulting in dominant form of dwarfism that results from a fail- coarseness of features, and is due to overproduction ure of cartilage to be converted into bone in the epi- of human growth hormone (hGH) secondary in the physeal disks. ACH affects mainly the long bones by majority of cases to an hGH-secreting pituitary

ad interim

adenoma (aka familial somatotrophinoma). This region of an enzyme or antigen-binding part of an condition is also obvious in skeletal remains; see antibody. paleopathology.Cf.gigantism. activity: 1. condition of being active. 2. release of acromelic shortening: congenital condition in which energy by nerve tissue. 3. intensity of a radioactive the most distal portions of a limb, i.e. the hands or element. the feet, are shortened. Cf. rhizomelic shortening, activity budget: amount of energy available for forag- mesomelic shortening. ing and feeding, movement, reproduction, and other acromial: pertaining to the shoulder region. activity beyond basal metabolism. Primates on a acromio-cristal index: bicristal breadth multiplied low-quality diet, such as folivores, are typically not by 100 and divided by the biacromial breadth. very active owing to the lack of energy for their activ- acromion: tip of shoulder that results from the lateral ity budgets. extension of the scapular spine. activity period: time of day when an animal is most acromion process: dorsal platelike extension of the active, especially in terms of foraging; e.g. diurnal, scapular spine that articulates with the clavicle; the nocturnal, crepuscular. acromion process provides sites for attachment of the actual extinction: physical extinguishment of a species muscles of the upper limb and chest. or germ line; actual extinctions occur during episodes acrosome: dense structure covering the anterior of mass extinction, or when a species is competitively half of the head of a spermatozoon that contains excluded from a niche, etc. See pseudoextinction. hyaluronidase, an enzyme that aids the penetration actualistic study: in archaeology, the study of the of the zona pellucida by the sperm during fertiliza- products and processes of present cultures as a key to tion. See capacitation. the past. Acrossia lovei Bown, 1979: anaptomorphine from acuity: sharpness, clarity, or distinctiveness, usually in middle Eocene of North America. Not recognized by reference to a sense. all authorities; some include Acrossia in Absarokius. acute: sudden; often severe, and short-lived; e.g. an Most notable features are enlarged central incisors acute illness. and relatively reduced size of the canines forming a acute mountain sickness: condition caused by scooplike arrangement suggestive of a gummivo- exposure to high altitude in which symptoms rous adaptation, although the cheek teeth are sug- appear in a few hours that include fatigue, dizziness, gestive of frugivory. See Appendix 1 for taxonomy. breathlessness, headache, nausea, vomiting, insom- ACTH: abbreviation for adrenocorticotropic hormone. nia, impairment of mental capacity and judgment, actinic ray: short-wavelength light that produces pho- and prostration. Not persistent; during acclimation tochemical effects, such as ultraviolet light. the body makes several physiological adjustments in actinomycosis: disease in humans characterized by breathing rate and red cell mass, and these symp- systemic soft tissue lesions and drainage sinuses, but toms abate. Cf. altitude sickness, chronic moun- that may progress to cartilage and bone. Affects tain sickness. males twice as often as females. Caused by two types acute promyelocytic leukemia (APL): malignant of bacillus, Actinomyces israelii and Arachnia propionica. proliferation of white blood cell precursors; see This condition is of interest to osteologists because it leukemia. APL is due to the abnormal transcription leaves its signature on the vertebral column, the of a fusion protein precipitated by a characteristic coxals, and bones of the hand. translocation of chromosomes 15 and 17 (t15;17) that action: 1. movement by the whole body. 2. particular causes a position effect when the gene for the effect of a speciﬁc skeletal muscle or muscle group retinoic acid receptor is located near an oncogene. when it contracts. 3. performance or function of any acute radiation: certain amount of radiation received organ or part. within a brief period. Cf. chronic radiation. active: capable of functioning or changing; requiring AD: 1. abbreviation for the (Latin) phrase anno Domini; energy. Cf. passive. since the birth of Christ. Used to indicate that a time active immunity: resistance to infection acquired division falls within the Christian era. Replaced by by contact with microorganisms, their toxins, or CE in some academic contexts. Cf. BC and BCE.2. other antigenic material such as by inoculation. abbreviation for autosomal dominant. Cf. acquired immunity. Ad Dabtiyah: archaeological site found in 1987 in active metabolism: any metabolic activity that gener- Saudi Arabia, faunal remains from which date to ates additional heat as a byproduct of the additional 17–16 mya. Hominoid remains include fragments of energy requirements of muscular work. Heliopithecus (cf. Afropithecus). active site: that portion of a protein that is required ad interim (ad. int.): (Latin) term used taxonomically for normal function, such as the substrate-binding to mean provisionally or temporarily; e.g. a new

adaptive

fossil may be classified Homo habilis ad int., meaning Adapis has a fused mandible. Estimated body size of that it appears that this fossil best fits into Homo the species around 1.5 kg. Small orbits suggest diur- habilis, but it may later be put into another or a new nality.Dental formula: 2.1.4.3/2.1.4.3; shearing crests species. on cheek teeth suggests folivory. Appears to have ad libitum (ad lib.): (Latin) at pleasure; applied in become extinct during the Grande Coupure (c. 38 science when no constraints are placed on the exper- mya). According to Gingerich, Adapis shows evidence imental subject. In ad libitum sampling there are no of sexual dimorphism; if so, this would be the set procedures as to what is recorded or when it is earliest appearance of this characteristic among the recorded. In primate behavior studies in which no primates. See Leptadapis and Appendix 1 for taxon- constraints are placed upon when or what is omy. recorded, the observer records any relevant behavior Adapoidea: superfamily of prosimians that consists that is visible during observation. This technique is of the families Adapidae, Notharctidae, and prone to bias in favor of conspicuous behavior or Sivaladapidae that are found from the Eocene to individuals. Aka haphazard sampling, convenience the Miocene. Adapoids are considered the most sampling. Cf. behavior, focal sampling, scan primitive euprimates. sampling. Adapoides troglodytes Beard, Qi, Dawson, and ADA: acronym for adenosine deaminase. Li, 1994: adapoid prosimian from the Eocene of ‡ Adapidae: family of prosimians known from the China assigned to the adapid subfamily Adapinae; Eocene and early Oligocene of Europe and Asia; sys- monotypic. Prior to the recovery of this fossil, tematics in flux, but currently about five adapid adapids were known only from Europe, although genera are recognized; origins and phylogenetic there is no evidence that they originated there; relationships uncertain. Adjectives: adapid, adapoid. the discovery of Adapoides suggests to some workers Earlier taxonomies included the notharctids that the adapids migrated to Europe from Asia. (Notharctidae)(as a subfamily) within the adapids, Body mass estimated at 500g. See Appendix 1 for which was a much larger group. See Adapinae and taxonomy. Appendix 1 for taxonomy. adaptation: 1. adjustment. 2. any alteration in struc- Adapiformes: infraorder of primates proposed by ture or function by which an organism becomes Szalay and Eric Delson that would include all of the better fitted to survive and reproduce in a given envi- lemur-like primates of the Eocene. ronment. See biological adaptation, behavioral Adapinae: subfamily of prosimians belonging to adaptation, functional adaptation, genetic adapta- the adapoid family Adapidae. Adjective: adapine. tion, insulative adaptation, metabolic adaptation, Known from the late Eocene to early Oligocene; five and sociocultural adaptation.Cf.preadaptation genera and twelve species. Recent revision of the and postadaptation. adapoids has resulted in this subfamily being adaptationist classification: proposed classification reduced in size as well as becoming the only subfam- of organisms using shared adaptations as data, ily of the adapids. All members are characterized by a rather than cladistic or phenetic similarity. Mayr molarized fourth premolar that is equipped with proposed the use of such a method to define para- shearing crests. Dental formula variable over time. phyletic higher taxonomic groups, such as birds, Unlike modern prosimians, the mandible is fused. which evolved rapidly from a common ancestor after Body mass estimated to range between 600g and achieving an adaptive breakthrough. 4 kg for the included genera. Analysis of the limb adaptationist program: perspective among some bones led some workers to suggest that the adapids evolutionary biologists in which all characters are employed a slow methodical climbing locomotion viewed as being optimized in adapting a species similar to extant lorisoids (Lorisoidea), but others to its environment. Researchers using this approach think that there was more diversity of locomotor attempt to determine the adaptive significance behavior in this group. See Appendix 1 for taxonomy. of species characters. The term was introduced in a Adapis Cuvier, 1821: well-known genus of primate discussion by Gould and Lewontin in 1979. See from the late Eocene to early Oligocene of western W. D. Hamilton,G.C. Williams,E.O. Wilson, and Europe and China belonging to the adapoid subfam- R. Dawkins. ily Adapinae; four to five species; first fossil primate adaptedness: state of the current fitness or func- described (by G. Cuvier, 1821), although its affinities tional superiority of one phenotype or genotype over were not recognized at the time. Possesses some another, as a result of past selection. lemurlike characteristics, which include an inflated adaptive: describes any trait that has arisen by the evo- auditory bulla with a free tympanic ring and some lutionary process of adaptation; the trait may be dental similarities; however, unlike modern lemurs, anatomical, physiological, or behavioral.

adeno-associated virus

Adaptive Behavior Scale: index of the range of economic factors that define human behavior from a adaptive behavior patterns measured in an organism research perspective. or society. adaptive trait: see adaptation and adaptive. adaptive breakthrough: evolution of a way of life adaptive value: reproductive success of an individual radically different from that of a recent common as measured by the mean value of its offspring with ancestor, which sometimes results in a perceived respect to a particular character. See fitness. paraphyletic higher taxonomic group, such as birds, adaptive zone: environment of previously unex- which then evolved rapidly from a common ancestor ploited ecological niches that becomes available to after achieving flight. The phrase could apply as well evolutionary opportunistic taxa; often results in an to certain hominids which, because of increased adaptive radiation. encephalization, are classified in the genus Homo additive: 1. effect of any element in a summation. 2. rather than as australopithecines. substance added to another material. 3. synergistic adaptive capacity: environmental tolerance of an effect. organism as determined by its genetic endowment. additive genetic effect: contribution an allele makes adaptive complex: suite of biological traits (includ- to the phenotype independent of the identity of ing behaviors) that enable a phylogenetic lineage to other alleles at the same or different loci. Aka addi- occupy a particular ecological niche. tive effect. See heritability.

adaptive evolution: see adaptation. additive genetic variation (Vga): proportion of the adaptive hormone: any substance secreted within the total phenotypic variance of a quantitative trait due body during adaptation to unusual circumstances, to genes that will actually be inherited in offspring. e.g. adrenocorticotropic hormone or the corticoids. See heritability. adaptive immunity: immune response that develops additive genotype fitness: hypothetical case where after exposure to a foreign antigen. fitness is the sum of a number of factors, such as the adaptive landscape: sinuous topographical graph of activities of the members of a family of digestive the average fitnesses of small, subdivided, and iso- enzymes. lated populations in relation to the frequencies of the additive hypothesis: in molecular taxonomy, a pro- genotypes residing in it. Peaks in such a landscape posal that parallelisms and convergences are ran- (multiple-peaked fitness surfaces) correspond to domly distributed among the taxa in any analysis. genotypic frequencies at which the average fitness is additive model: model of heritability in which the high; valleys to genotypic frequencies at which the total phenotypic variance of a quantitative trait is average fitness is low. Proposed by Sewall Wright. partitioned into several genetic and environmental Aka adaptive topography, fitness surface, surface of subcomponents. selective value. See shifting balance theory. additive technology: processes in which manufac- adaptive mechanism: any device, idea or process that tured artifacts take form by the addition of materials increases an organism’s chances for survival, e.g. to the original mass, such as ceramic production or mechanisms of culture change and mechanisms basket making. Cf. subtractive technology. of evolution. adductor: muscle that moves a limb or structure adaptive niche: see ecological niche. toward the mid-line; aka depressor. Cf. abductor. adaptive prediction: theoretical relationship between adductor pollicis: intrinsic muscle of the hand; certain morphs and environmental variables; if a trait originates from the trapezium, trapezoid, capitate, is an adaptation to an external condition, then the and metacarpals II–IV, and inserts into the medial value of that condition should predict morphology. surface of proximal phalanx I. Its action is to adduct Bergmann’s rule, predicts, for example, that organ- and oppose the thumb; one of the thenar group of isms with adaptations for high average annual tem- muscles. perature should be found nearer to the equator than ‡ adenine (A): one of the purine nitrogenous bases organisms with adaptations to low temperatures. that composes DNA and RNA; composed of two ‡ adaptive radiation: biologic evolution in a group of carbon–nitrogen rings. Adenine bonds with thymine related species that is characterized by spreading into in DNA and with uracil in RNA (see base pairing different environments and by divergence of struc- rule); it is also a major component of other molecules ture, e.g. the 14 species of finch in the Galápagos such as adenosine triphosphate. archipelago, and hundreds of Hawaiian fruit fly adeno-associated virus (AAV): any one of a group of species (the ‘Hawaiian radiation’). defective spherical DNA viruses frequently isolated adaptive strategy: 1. totality of mechanisms pos- from tonsillar tissue that can replicate only in the sessed by a taxon that help it survive in an environ- presence of adenoviruses. No disease has been asso- ment. 2. technological, ecological, demographic and ciated with AAV in humans. When not replicating it

adrenal

is integrated into the host chromosome. AAV is most adiacritic: pertaining to the condition in which no notable as a proposed vector for human gene transfer member of a race or breed is recognizable as such experiments. by an expert. Cf. typology, microdiacritic,and adenohypophysis: anterior portion of the pitu- mesodiacritic. itary gland, and which secretes most of its hor- adipocyte: fat cell. mones, including growth hormone. See pituitary adipometer: anthropometric instrument used for gonadotropic hormones.Cf.neurohyphophysis. measuring skinfold thickness in order to estimate adenoma: tumor or growth located in glandular the amount of subcutaneous fat. tissue. adipose tissue: specialized connective tissue that adenomatous polyposis of the colon, familial stores fat; in humans much of this tissue is found (FAP): one of the two most common forms of heredi- in the subcutaneous layer of the skin. It is also tary colorectal cancer. FAP is an autosomal dominant found in spaces between muscles, behind the eyes, found in adolescents who present with abdominal around the kidneys, in some abdominal mem- pain, diarrhea and rectal bleeding. A carpet of pre- branes, on the surface of the heart, and around cer- cancerous growths, usually in the colon, progresses tain joints. to colorectal cancer. The mutant gene is adenoma- adiposis: corpulence or obesity; an excessive accumu- tous polyposis coli (APC); about 740 different muta- lation of body fat. tions in this gene have been identified with adiposity: amount of body fat; state of being fat. Often simultaneous loss of heterozygosity or mutation used in reference to excessive obesity, esp. in medi- to the p53 tumor suppressor gene. There is some cine. evidence for a paternal age effect. Aka familial poly- adjustment: physiological or behavioral response of posis of the colon (FPC). Cf. colon cancer, heredi- an organism to change in its environment, without tary nonpolyposis. regard to whether the change is beneficial to the adenosine deaminase (ADA): an enzyme that con- organism. See accommodation. verts adenosine to inosine in the purine salvage path- admixture: term used for interbreeding between races; way, and is essential to the production of T cells and in humans, miscegenation; in evolutionary terms, B cells in the immune system. The ADA gene is gene flow. located on HSA 20q13.1. Some 95% of the ADA adolescence: another term for the adolescent stage. cistron consists of introns; the 12 exons have a final adolescent: see adolescence.Cf.juvenile and adult. translation length of 1500 base pairs. Cf. purine ‡ adolescent growth spurt (AGS): a rapid increase in nucleoside phosphorylase. stature during adolescence. It occurs at the same adenosine deaminase deficiency: an autosomal developmental stage in all primates regardless of the dominant condition, one of the primary immuno- age of the individual in years. See hypertrophy, deficiency diseases (PIDs) characterized by skeletal hyperplasia and life-history variable. and neurological abnormalities; frequent infections, ‡ adolescent stage (or period): interval in the fatal if untreated. Death usually occurs by the age of human life cycle preceeding adulthood that is char- 7 months owing to infection. ADA deficiency acterized by the adolescent growth spurt in height accounts for about 15% of all severe combined and body mass, fusion of the epiphyses of the long immune deficiency (SCID) cases. There are no T cells, bones, development of secondary sexual characteris- and B cells do not produce antibodies. An ADA- tics, continuation or completion of adult tooth erup- deficient SCID girl was selected as the first person tion, sociosexual maturation, and intensification of with a genetic disorder to be treated by somatic gene interest in and practice of adult social roles. The stage therapy, using a viral vector, in September 1990. Cf. commences at puberty and lasts five to eight years; agammaglobulinemia. aka adolescence. adenosine triphosphate: see ATP. adolescent sterility: physiological state in adolescent adenovirus (AV): any member of a group of spherical females that begins with menarche and ends with the DNA viruses that infect several mammalian species, onset of regular ovulatory cycles. including humans; some are oncogenic. adontia: lack of tooth bud development; see ectoder- adenylate kinase (AK): an enzyme that releases mal dysplasia. See also hyperdontia, hypodontia. energy within cells; in humans, two common alleles adoption: shared welfare of offspring by individuals are known, AK-1 and AK-2. AK is of historical other than biological parents; a behavior found in significance because it is linked to both the ABO many species. Contrasts with strict biological or blood group locus and the Nail Patella locus; this genetic modes of defining human relationships. linkage group is one of the first discovered, now adrenal: 1. located near the kidney. 2. pertaining to the known to reside on chromosome 9q. adrenal gland; adrenogenic.

adulthood

adrenal cortex: outer layer of the adrenal gland; inevitable by age 3. Variable symptoms: the X-linked secretes mainly cortisol and other glucocorticoids, form is less severe than the neonatal autosomal reces- aldosterone, and androgens. sive form. The defect is in the ALD membrane trans- adrenal gland: paired endocrine gland located super- port protein; adrenal insufficiency causes an excess of ior to each kidney in most vertebrates; a composite long-chain fatty acids. Maps to Xq 28. The condition organ that consists of the adrenal medulla and the is associated with HLA DR3 and increases the relative adrenal cortex of differing embryonic origin that risk of ALD to 6.3ϫ. ALD is rare, incidence 1:100 000. function to secrete several ‘stress’ hormones. Aka Aka Addison disease. suprarenal gland, epinephric gland, third gonad. adrenomegaly: enlargement of the adrenal glands. adrenal hypoplasia, primary: heritable disease char- adrenosterone: androgenic steroid present in the acterized by delay or absence of puberty, possible adrenal cortex. See androgen. mental retardation, and possible death at puberty adrenotropin: 1. see adrenocorticotropic hormone. without hormonal supplementation. Caused by 2. see corticotropin. Also spelled adrenotrophin. mutations in genes that disrupt the adrenal glands adulation: transfer of the appearance of a trait from a and by hormone deficiencies. late to an earlier stage of development. Cf. neoteny. adrenaline: alternative name for the hormone and adult: in mammals, including humans, an individual neurotransmitter epinephrine; produced by the that has reached full growth in height or length; note adrenal medulla of the kidney. Adrenaline initiates that reproductive adulthood can occur before full actions similar to those initiated by sympathetic body size is reached. Cf. adolescent. nervous system stimulation (andrenergic stimula- adult body size: mass or dimension of an organism tion) in the heart, bronchioles and blood vessels. during maturity. Cf. body size. adrenal medulla: soft center of the adrenal gland; adult dentition: set of teeth in mammals that replaces secretes mainly epinephrine and norepinephrine. the deciduous dentition; in most mammals the adrenarche: period when secretion of the adrenal adult set consists of incisors, canines, premolars, and androgen hormones begins; has been suggested to be molars. The larger adult dentition fits the growing involved with the mid-growth spurt that occurs in adolescent, and later adult, jaw. Aka permanent children between the ages of six and eight. See teeth, replacement dentition, adult teeth. androgen and mid-growth spurt. adult form: in genetics and physiology, the form of a adrenergic: pertaining to nerve fibers of the sympa- molecule that is present in the adult, as opposed thetic nervous system that, upon stimulation, to forms present in the embryo, fetus, or other release the chemical transmitter norepinephrine developmental stage. The hemoglobin molecule, for (and possibly epinephrine) at their post-ganglionic example, is found in three forms during the corres- endings; any compound that acts like norepineph- ponding stages of development: embryonic, fetal, rine or epinephrine. and adult hemoglobin. adrenocortical hormones: steroids produced by the adult hemoglobin: umbrella term for three forms adrenal cortex, including estrogens, androgens, of hemoglobin found in the erythrocytes of progesterone, and the glucocorticoids. adult humans that are metabolically active in the adrenocorticotropic hormone (ACTH): a polypep- late fetal stages of development through adult- tide secreted by the anterior pituitary gland; it con- hood. The majority consists of hemoglobin A (aka

trols secretion of certain hormones called adrenal the major fraction) and about 2% is hemoglobin A2, androgens (e.g. cortisol) that originate in the the minor fraction; a very small percentage consists adrenal cortex, and assist in the breakdown of fats. of the so-called fetal fraction, which, however, is Aka adrenocorticotropin, corticotropin. not quite identical to fetal hemoglobin in that the adrenocorticotropin: see adrenocorticotropic hor- fraction circulating in normal adults has an amino mone. acid substitution at position 136 in the ␥ chain com- adrenogenital syndrome: condition resulting from pared with the normal fetal form. Cf. embryonic a hereditary defect in the adrenal glands in which a hemoglobin. nonfunctioning enzyme in a metabolic pathway is adult polycystic kidney disease: see polycystic produced, causing the accumulation of testosterone- kidney disease, adult. like breakdown products that cause affected females adulthood: interval of maturity that commences at to exhibit masculine characteristics and to develop about age 20 years. The stage in the human life cycle rudimentary male sex organs. between adolescence and senescence; the prime adrenoleukodystrophy (ALD): characterized by of adulthood lasts until the end of child-bearing dementia, seizures, paralysis, loss of speech, deaf- years and is a period of homeostasis in physiology, ness, and blindness; neonatal death is usual, and behavior and cognition. Cf. middle age.

‡ affiliative

adult teeth: see adult dentition. aetiology: see etiology. advanced: see derived. Afalou man: AMH fossil found in North Africa, and advanced age: state of living into post-reproductive similar to Cro-Magnon. life; living after the majority of one’s birth cohort has Afalou-Bou-Rhommel: Upper Pleistocene rockshel- already expired. See maturity, senility. ter found in 1928 near Bugia in Algeria, dated to advanced parental age effect(s): increase in the 15–8kya, that contains artifacts identified as frequency of a human trait when either the mother is Oranian (ϭ Capsian) and closely related to over about 35 years of age (see Down syndrome and Aurignacian. Hominid remains include at least 50 dizygotic twins), or the father is over about 50 years skeletons comparable in morphology to Cro- of age (see Marfan syndrome and other collagen Magnon in Europe (i.e. Homo sapiens). All the diseases). See maternal age effect and paternal individuals had experienced dental mutilation. Aka age effect. Afalou-Bou-Rhummel, Mechta-el-Arbi. The arti- adventive: in reference to an organism not native to a facts are aka Ibero-Maurusien. particular region. Aka exotic; alien; introduced. Afar depression: region in Ethiopia; the west central Aedes: genus of mosquito (family Culicidae) that con- Afar sedimentary basin contains several hominid- tains over 700 species and that is a potential vector of bearing sites (e.g. Hadar, Belohdelie, Maka) that yellow fever and malaria; e.g. Aedes aegypti. date to earlier than 2.9 mya, and possibly as far back ‡ Aegyptopithecus Simons, 1965: extinct genus of as 3.6 mya. largest and best known of the haplorhines of the Afar hominids: see entries beginning with NME. Oligocene epoch (c. 34–33 mya); recovered from the Afar Locality (AL): any site located in the Afar Fayum Depression, Egypt; member of family Depression of Ethiopia. Thus AL-288 is the specifi- Propliopithecidae. Completely anthropoid in char- cation for the site where the australopithecine acteristics such as closed orbits, fusion of frontal known as ‘Lucy’ was found (field number NME and mandibular bones. Highly sexually dimorphic; AL-288-1) in 1974, NME AL-333 where the ‘first sagittal crest and long canines present in males. family’ was found in 1975, and so forth. Small orbits indicate diurnal habits. Brain size about Afar triangle or Afar region: see Afar depression. 30 cm3, and comparable with that of extant prosimi- afarensis nomen debate: difference of opinions con- ans. Elongated maxilla; dental formula: 2.1.2.3; denti- cerning the attribution of fossils found at Laetoli and tion suggests frugivory. Estimated body mass 6–8 kg. Hadar; one group (led by Mary Leakey) felt that Monotypic; the only known species is A. zeuxis,which these early pre-habilines should be placed in the has affinities with both monkeys and apes; it could be genus Homo, while another (Johanson, White, and ancestral to one or both of these groups. Some author- others) erected the taxon Australopithecus afaren- ities now include Aegyptopithecus in Propliopithecus. sis for these specimens. The first group felt that the See Appendix 1 for taxonomy. new taxon contained more than one species and that, aeolian: alternative spelling of eolian. as the holotype had come from Laetoli and not the Aeolopithecus Simons, 1965: genus of the fossil pri- Afar triangle, the selection of the specific nomen was mate family Propliopithecidae, an anthropoid from itself inappropriate. Tobias had suggested sub- the Oligocene epoch. Synonym: Propliopithecus, specific taxa to resolve the issue (A. africanus aethiopi- which is accepted now by most authorities. Recent cus for Johanson’s Ethiopian fossils, and A. A. considerations of sexual dimorphism within P.chiro- tanzaniensis for Leakey’s), but Johanson presented his bates has removed this genus as a valid taxon. See description first, thus establishing priority. Propliopithecus; See Appendix 1 for taxonomy. affection: expression of emotions, feelings or mood aerobic capacity: highest rate at which an indi- indicating tenderness, one of the emotions mediated vidual is capable of utilizing oxygen. See aerobic by the limbic system. endurance. affective disorder: any of a group of similar behav- aerobic endurance: length of time that an organ, ioral disorders pertaining to feelings; alternation particularly muscle tissue, can continue to use oxygen between manic and depressive behavior is called in metabolic pathways. See aerobic capacity. bipolar affective disorder, whereas the experience aerobic power: aerobic capacity divided by body of depressive bouts alone is called unipolar disorder. mass. See psychiatric disorder. aestivation: see estivation. afferent: towards a reference point; for example, affer- Aethiopian: early ethnic term, dating from antiquity, ent nerves conduct impulses towards the central meaning ‘person with a burnt face’. Used by Homer, nervous system. Xenophanes, Scylax of Caryanda, and Herodotus. See ‡ affiliative: pertaining to persistent amicable rela- Ethiopian. tions between individuals.