Showing papers in "Systematic Biology in 1980"

Non-Allopatric Speciation in Animals

Abstract: Futuyma, D. J., and G. C. Mayer (Department of Ecology and Evolution, State University of New York, Stony Brook, New York 11794 and Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138) 1980. Non-allopatric speciation in animals. Syst. Zool., 29:254-271.-Major recent challenges to the view that animal speciation is usually allopatric are reviewed, and are found unconvincing, either because of their theoretical implausibility or because of insufficient evidence. Special attention is given to the theory of stasipatric speciation, and to purported cases, especially in tephritid fruit flies, of sympatric speciation associated with a shift to a new host. Stasipatric speciation is unlikely under population genetic theory; moreover, chromosome rearrangements probably seldom facilitate speciation. In the tephritid genus Rhagoletis, the archetypal case of sympatric speciation, there is little or no evidence of genetic divergence or of sympatric speciation. The conditions under which host-associated sympatric speciation might occur are so exacting as to be met by very few species. [Rhagoletis; Didymuria; Vandiemenella; Mus; Sceloporus; speciation; sympatric speciation; parapatric speciation; allopatric speciation; chromosome races; isolating mechanisms; host selection.] "It is in the nature of science that once a position becomes orthodox it should be subjected to criticism .... It does not follow that, because a position is orthodox, it is wrong .... -Maynard Smith, 1976 Until recently, it has been widely accepted that speciation in animals usually proceeds by the differentiation of geographically isolated populations (e.g., Dobzhansky, 1970; Lewontin, 1974; Rosenzweig, 1975; Dobzhansky et al., 1977). The reasons for this belief were developed in extenso by Mayr (1942, 1963, 1970), who argued that other modes of speciation were so unlikely as to be insignificant. Since 1963, there have been several important challenges to this view (e.g., Maynard Smith, 1966; Bush, 1975a; White, 1978a). Because White's (1978a) book, devoted largely to demonstrating the prevalence of non-allopatric speciation, seems likely to have an important impact on the climate of opinion on this topic, it is useful at this time to review the extent to which the theory and evidence amassed since 1963 warrant a major change in our views of animal spe-

Significance Tests for Coefficients of Variation and Variability Profiles

Abstract: Sokal, R. R., and C. A. Braumann (Department of Ecology and Evolution, State University of New York at Stony Brook, Stony Brook, New York 11794) 1980. Significance tests for coefficients of variation and variability profiles. Syst. Zool. 29:50-66.-The distribution of sample estimates of the coefficient of variation is studied analytically and by Monte Carlo simulation. Derivations are given for the expected value of a coefficient of variation and for its standard error. Various proposed standard errors for coefficients of variation are evaluated. Standard errors are derived for differences between coefficients of variation for samples of independent and correlated characters. Methods are proposed for testing the homogeneity of sets of independent and correlated coefficients of variation. Tests of homogeneity of variability profiles as well as for parallelism of such profiles are furnished. [Coefficients of variation, variability profiles.] The employment of coefficients of variation in systematic research is of long standing. Various evolutionary hypotheses require for their examination the establishment of differences in the amounts by which characters vary in populations. Such differences can be examined for the same character across several populations of the same species or of different species, or the comparison may be within the same population but among different characters. Such comparisons of the amounts of variation are generally adjusted for differences in magnitude of the character means, hence the employment of the coefficient of variation, V. A recent renewal of interest in the coefficient of variation is due to two developments. Various studies, collectively called population phenetics, have probed the effects of evolutionary processes on variability patterns in animal and plant populations and have tried to establish the converse-the drawing of inferences about evolutionary processes from observed variability patterns. Studies such as those of Soule (1967), Soule and Stewart (1970), Rothstein (1973), Lande (1977), or Sokal (1976) come to mind readily. A second reason for an increased interest in coefficients of variation is the stimulating book by Yablokov (1974) introducing the study of variability profiles in mammalian populations. Variability profiles are graphs in which the amount of variation expressed as a variance or coefficient of variation is plotted against a horizontal axis representing the suite of characters under study. Examination of variability profiles within and among populations leads to inferences about the amount of developmental (and ultimately evolutionary) control of variability for different characters in the same population and among populations. There is a need for appropriate methods to examine the types of comparisons being considered. In this paper we shall briefly review the expectations and variances of coefficients of variation under the assumption of normality and examine the effects of appreciable departures from this assumption. We shall then turn to the comparison of two or more coefficients of variation for the same character from different populations. This account will be followed by a discussion of tests applicable to a single variability profile, which in turn will lead to the comparison of several profiles. These can be considered for the same hierarchic level, as in local population samples of the same species, or the comparison may be between different hierarchic levels representing natural sampling units, such as variability within local populations versus variability across populations. Analytical work on the expectations

Comparative biology and evolutionary relationships of tree shrews

Abstract: s of Papers, Jour. Paleont. 51 (suppl. 2, part III): 10. 90 MICHAEL]. NOVACEK Englemann, G. F. and E. O. Wiley. 1977. The place of ancestor-descendant relationships in phylogeny reconstruction. Syst. Zool. 26: 1-12. Evans, F. G. 1942. Osteology and relationships of the elephant shrews (Macroscelididae). Bull. Amer. Mus. Nat. Hist. 80: 85-125. Frey. H. 1923. Untersuchungen tiber den Scapula, speziell tiber aussere Form und Abhangigkeit von Funktion. Zeitschr. Gesamt. Anat. 68: 276-324. Fuchs, H. 1910. Uber das Pterygoid, Palatinum, und Parasphenoid der Quadrupeden, insbesondere der Reptilia und Saugetiere. Anat. Anz. 36:33-95. Gamberian, P. P. and R. O. Oganesian. 1970. Biomechanics of the gallop and the primitive rebounding jump in small mammals. Proc. Acad. Sci. U.S.S.R. Bioi. Ser. 3: 441-447. Gingerich, P. D. 1973. Anatomy of the temporal bone in the Oligocene anthropoid Apidium and the origin of the Anthropoidea. Folia Primat. 19:329-337. Gingerich, P. D. 1974. Stratigraphic record of early Eocene Hyopsodus and the geometry of mammalian phylogeny. Nature 248: 107-109. Gingerich, P. D. 1976. Cranial anatomy and evolution of the early Tertiary Plesiadapidae (Mammalia, Primates). Mus. Paleont. Univ. Michigan Papers on Paleontology 15: 1-117. Goodman, M. 1975. Protein sequence and immunological specificity: Their role in phylogenetic studies of primates, pp. 219-248. In W. P. Luckett and F. S. Szalay (eds.). Phylogeny of the Primates. Plenum Press, New York. Gregory, W. K. 1910. The orders of mammals. Bull. Amer. Mus. Nat. Hist. 27: 1-524. Gregory, W. K. 1920. Studies of the comparative myology and osteology; no. IV A review of the evolution of the lacrimal in vertebrates with special reference to that of mammals. Bull. Amer. Mus. Nat. Hist. 42: 95-263. Haeckel, E. 1866. Generelle Morphologie der Organismen. G. Reimer, Berlin. Haines, R. W. 1950. The interorbital septum in mammals.Jour. Linnean Soc. London (Zool.) 41: 585607. Hall-Craggs, E. C. B. 1965. An analysis of the jump of the lesser galago (Galago senegalensis). Proc. Zool. Soc. London 147: 20-29. Hatt, R. T. 1932. The vertebral column of richochetal rodents. Bull. Amer. Mus. Nat. Hist. 63: 599738. Hecht, M. K. 1976. Phylogenetic inference and methodology as applied to the vertebrate record. Evol. Bioi. 9: 335-363. Hecht, M. K. and j. Edwards. 1976. The determination of parallel or monophyletic relationships: The proteid salamanders-a test case. Am. Nat. 110: 653-677. Hennig, W. 1950. Grundzuge einer Theorie der Phylogenetischen Systematik. Deutscher Zentral Verlag, Berlin. Hennig, W. 1966. Phylogenetic Systematics. University of Illinois Press, Urbana. Hennig, W. 1975. ·Cladistic analysis or cladistic classification?\": A reply to Ernst Mayr. Syst. Zoo!. 24: 244-256. Hildebrand, M. 1974. Analysis of Vertebrate Structure. john Wiley and Sons, New York. Howell, A. B. 1932. The saltatorial rodent Dipodomys, the functional and comparative anatomy of its muscular and osseous systems. Proc. Am. Acad. Arts Sci. 67: 377-536. Howell, A. B. 1944. Speed in Animals, Their Specializations for Running and Leaping. Hafner, New York. Hull, D. L. 1970. Contemporary systematic philosophies. Ann. Rev. Eco!. Syst. I: 19-54. Hunt, R. M., jr. 1974. The auditory bulla in Carnivora: An anatomical basis for reappraisal of carnivore evolution. Jour. Morph. 143: 21-76. jenkins, F. A.,jr. 1971. Limb posture and locomotion in the Virginia opossum (Didelphis marsupialis) and other non-cursorial mammals. Jour. Zoo!.. 165: 303-315. jenkins, F. A., jr. 1974. Tree shrew locomotion and the origins of primate arborealism, pp. 85115. In F. A. jenkins, jr. (ed.) Primate Locomotion. Academic Press, New York. jepsen, G. L. 1970. Bat origins and evolution, pp. 1-64. In W. A.Wimsatt (ed.) Biology of the Bats. Volume I. Academic Press, New York. j ouffray, F. K. 1975. Osteology and myology of the lemuriform postcranial skeleton, pp. 149-192. In I. Tattersall and R.W. Sussman (eds.). Lemur Biology. Plenum Press, New York. CRANIOSKELETAL FEATURES IN TUPAIIDS 91 Kavanaugh, D. H. 1972. Hennig's principles and methods of phylogenetic systematics. The Biologist 54: 115-127. Kay, R. F. and M. Cartmill. 1974. The skull of Palaechthon nacimienti. Nature 252: 37-38. Kermack, K. A. and Z. Kielan-Jaworowska. 1971. Therian and non-therian mammals. Zool. Jour. Linnean Soc. London (suppl. 1): 103-115. Kielan-Jaworowska, Z. 1977. Evolution of therian mammals in the Late Cretaceous of Asia. Part II. Postcranial skeleton in Kennalestes and Asioryctes. Palaeont. Polonica 37: 65-83. Krishtalka, L. 1976. Early Tertiary Adapisoricidae and Erinaceidae (Mammalia, Insectivora) of North America. Bull. Carnegie Mus. Nat. Hist. I: 1-40. Kruger, W. 1958. Der Bewegungsapparat, pp. 1-176.In W. Kukenthal (ed.). Handbuch der Zoologie, Vol. 8 (13-14). Walter de Gruyter and Co., Berlin. Le Gros Clark, W. E. 1934. On the skull structure of Pronycticebus gaudryi. Proc. Zool. Soc. London 1934: 19-27. Le Gros Clark, W. E. 1959. The Antecedents of Man. 1st ed. Edinburgh University Press, Edinburgh. Le Gros Clark, W. E. 1971. The Antecedentl of Man. 3rd ed. Quadrangle Books Inc., Chicago. Lessertisseur,J. and R. Saban. 1967. Squelette appendicular. In P. P. Grasse (ed.) Traite de Zoologie; Anatomie, Systematique Biologie. Mammifires, Tegumentl Squelette. Tome XVI, Fasc. I. Masson et Ce, Paris. Llorca, F. O. 1934. L'artere stapedienne chez l'embryon humain et la cause mecanique probable de son atrophie. Arch. Anat. Antrop. Lisboa 16: 199-207. Luckett, W. P. 1969. Evidence for the phylogenetic relationships of tree shrews (family Tupaiidae) based on the placenta and foetal membranes.]. Reprod. Fert., Suppl. 6: 419-433. MacIntyre, G. T. 1972. The trisulcate petrosal pattern of mammals. Evolut. Bioi., 6: 275-303. MacPhee, R. D. E. 1979. Entotympanics, ontogeny and primates. Folia Primat. 31:23-47. McDowell, S. B., Jr. 1958. The Greater Antillean insectivores. Bull. Am. Mus. Nat. Hist. ll5: ll3214. McKenna, M. C. 1966. Paleontology and the origin of the primates. Folia Primat. 4: 1-25. McKenna, M. C. 1975. Toward a phylogenetic classification of the Mammalia, pp. 21-46. In W.P. Luckett and F. S. Szalay (eds.). Phylogeny of the Primates. Plenum Press, New York. Matthew, W. D. 1909. The Carnivora and Insectivora of the Bridger Basin, middle Eocene. Mem. Am. Mus. Nat. Hist. 9: 291-567. Matthew, W. D. 1937. Paleocene faunas of San Juan Basin, New Mexico. Trans. Am. Philos. Soc. N. S. 30: 1-510. Mayr, E. 1974. Cladistic analysis or cladistic classification? Z. Zool. Syst. Evol.-Forsch. 12: 94-128. Miller, G. S., Jr. 1907. The families and genera of bats. Bull. U. S. Nat. Mus. 57: 1-282. Muller, J. 1934. The orbitotemporal region in the skull of the Mammalia. Archiv. Neerl. Zool. I: 118-259. Nelson, G. J. 1973. The higher level phylogeny of the vertebrates. Syst. Zool. 22: 87-92. Novacek, M. J. 1976. Insectivora and Proteutheria of the later Eocene (Uintan) of San Diego County, California. Nat. Hist. Mus. Los Angeles County Contr. Sci. 283: I-52. Novacek, M. J. 1977a. A review of Paleocene and Eocene Leptictidae (Eutheria: Mammalia) from North America. PaleoBios 24: 1-42. Novacek, M. J. 1977b. Evolution and relationships of the Leptictidae (Eutheria: Mammalia). Ph. D. Thesis, University of California, Berkeley. Novacek, M. J. 1977c. Aspects of the problem of variation, origin, and evolution of the eutherian auditory bulla. Mammal Rev. 7: 131-149. Patterson, B. 1965. The fossil elephant shrews (family Macroscelididae). Bull. Mus. Compo Zool. 133: 295-385. Piveteau, J. 1957. Traite de Paleontologie, Vol. 7. Masson et CeParis. Platnick, N. I. 1977. Parallelism in phylogeny reconstruction. Syst. Zool. 26: 93-96. Roberts, D. and I. Davidson. 1975. The lemur scapula, pp. 125-147. In I. Tattersall and R. W. Sussman (eds.). Lemur Biology. Plenum Press, New York. Rose, K. D. and E. L. Simons. 1977. Dental function in the Plagiomenidae: Origin and relationships of the mammalian order Dermoptera. Contr. Mus.Paleon. Univ. Michigan 24: 221-236. 92 MICHAEL J. NOVACEK Saban, R. 1963. Contribution a l'etude de l'os temporal des primates. Mem. Mus. Natl. d'Hist. Nat. (nouv. ser., ser. A.) 29: 1-378. Salomon, M. I. 1930. Considerations sur l'homologie de l'os lachrymal chez les Vertebres superieurs. Acta Zool. 11: 151-183. Schaeffer, B., M. K. Hecht, and N. Eldredge. 1972. Phylogeny and paleontology. Evol. Bioi. 6: 3146. Sige. B. 1974. Pseudorhynchocyon cayluxi Filhol, 1892, insectivore geant des phosphorites du Quercy. Palaeovertebrata 6: 33-46. Simons, E. L. 1962. A new Eocene primate genus, Cantius;'and a revision of some allied European lemuroids. Bull. Brit. Mus. Nat. Hist. Geol. 7:1-36. Simons, E. L. 1972. Primate Evolution, An Introduction to Man's Place in Nature. MacMillan, New York. Simpson, G. G. 1945. The principles of classification and a classification of mammals. Bull. Am. Mus. Nat. Hist. 85: 1-350. Simpson, G. G. 1975. Recent advances in methods of phylogenetic inference, pp. 3-19. In W. P. Luckett and F. S. Szalay (eds.). Phylogeny of the Primates. Plenum Press, New York. Sokal, R. R. 1975. Mayr on cladism-and his critics. Syst. Zool. 24: 257-262. Spatz, W. B. 1966. Zur Ontogenese der Bulla Tympanica von Tupaia glis Diard 1820 (Prosimiae, Tupaiiformes). Folia Primat. 4: 26-50. Szalay, F. S. 1968. The beginnings of primates. Evolution 22: 19-36. Szalay, F. S. 1969. Mixodectidae, Microsyopidae and the insectivore-primate transition. Bull. Am. Mus. Nat. Hist. 140: 197-330. Szalay, F. S. 1972. Cranial morphology of the early Tertiary Phenacolemur and its bearing on primate phylogeny. Am. J. Phys. Anthrop. 36: 59-76. Szalay, F. S. 1975. Phylogeny of primate higher taxa: The basicranial evidence, pp. 91-125. In W. P. Luckett and F. S. Szalay (eds.). Phylogeny of the Primates. Plenum Press, New York. Szalay, F. S. 1977a. Ancestors, descendants, sister groups and testing of phylogenetic hypotheses. Syst. Zool. 26: 12-19. Szalay, F. S. 1977b. Phylogenetic

Karyotypic Evolution in Bats: Evidence of Extensive and Conservative Chromosomal Evolution in Closely Related Taxa

Abstract: Baker, R.J. , and] . W . Bickha,m (The Museum and Department of Biological Sciences, Texas Tech University, Lubbock, Texas 79409 and Department of Wildlife and Fisheries Sciences, Texas A and M University, College Station, Texas 77843) 1980. Karyotypic evolution in bats: evidence of extensive and conservative chromosomal evolution in closely related taxa. Syst. Zool., 29:239-253.-Gand C-band data for seventy-eight species of bats from four families were subjected to a cladistic analysis to determine the number of chromosomal rearrangements required to convert the karyotype proposed as primitive for a family into the karyotype of extant species in that family. The number of rearrangements ranged from 0 to 36, and if the age of families is 60 million years, average rate of incorporation of rearrangements per million years ranged from 0 to 0.6. When chromosomal variation in congeneric species were subjected to a similar-dadistic analysis, most (34 of 54) species had undergone no chromosomal rearrangements; however, some species had undergone from 14 to 20 rearrangements and the types of rearrangements that were incorporated in species having the largest amount of change were generally rearrangements that should produce considerable reduction in gamete fertility in individuals heterozygous for such rearrangements. Radically reorganized karyotypes appear not only in bats but in a wide variety of vertebrates. Factors related to demography, breeding structure, and speciation do not appear adequate to explain the occurrence of such radically reorganized genomes. Factors less related to demographic and vagility characteristics, such as mutation rate and mechanisms which reduce the meiotic constraints on the heterozygote, are phenomena which may be involved in evolving a radically reorganized karyotype. [Chromosomal evolution; G-bands; rates of evolution; cladistic analysis; bats.] In order to understand the meaning of Also, we describe and discuss a phenomchromosomal evolution. it is necessarv to enon which we call "karyotypic megahave accurate estimates of magnitude of evolution" which appears to occur not chromosomal change and documentation only in bats but in a wide variety of living of the degree to which the evolution of forms. species with differing biological characteristics has been associated with karyoMETHODS AND MATERIALS typic change. Knowledge of rates are imAll species of bats included in this reportant in the development and testing of port have been studied from Gand models of chromosomal evolution (Bush C-band preparations. Most of these e t al., 1977; Bickham and Baker, 1979; karyotypes have been described elseLande, 1979; Bengtsson, 1980). Th' is rewhere (Bickham and Baker, 1977, table port is concerned with results of G-band 1; Bickham and Hafner, 1978; Bickham, studies of the bat families Phyllostomat1979a and 197913; Bass, 1978; Baker, idae (37 species), Vespertilionidae (29 1979; Baker et al., 1979a; Johnson, 1979; species), Mormoopidae (6 species), and Patton and Baker, 1978). Noctilionidae (2 species). Approximately Within each family, chromosomal data nine percent of known bat species are inwere arranged such as to require the cluded in this r e ~ o r t . We give estimates minimum number of events derive to of magnitude of change in lineages exkaryotypes of living species. From such tending back to the hypothesized primiarrangements, it was possible to hypothtive karyotype for each family and shorter esize a primitive karyotype for each famterm estimates from congeneric species. ily (and in the case of the Phyllostomat-

Analysis of taxonomic congruence among morphological, ecological, and biogeographic data sets for the Leptopodomorpha (Hemiptera)

Abstract: Schuh, R. T., andJ. T. Polhemus (Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024 and 3115 S. York, Englewood, Colorado 80110) 1980. Analysis of taxonomic congruence among morphological, ecological and biogeographic data sets for the Leptopodomorpha (Hemiptera). Syst. Zool. 29:1-26.-A data set of 47 morphological characters derived from the literature and original observation is prepared for the Leptopodomorpha. An argument is presented for considering the Leptopodomorpha as a monophyletic group and the Nepomorpha as its sister group. The data set is analyzed by cladistic (Wagner) and phenetic (UPGMA) methods in an effort to arrive at a most natural classification. Predictivity and stability are measured by comparing classifications based on a complete and partial taxon set as well as a random bipartition of the character set. Cladistic analysis produces more stable and predictive classifications with greater consensus and component information. The minimum length Wagner tree has greater information content than the UPGMA phenogram by virtue of its higher cophenetic correlation coefficient and its more parsimonious description of the character data. The Wagner tree is compared with four existing classifications for the Leptopodomorpha in the form of networks and trees. One of the published schemes is represented by the same network as the most parsimonious tree, but contains additional homoplasies as a rooted tree. The remaining published schemes are represented by different networks and describe the data less accurately (in more steps) than the most parsimonious tree. Wagner, UPGMA, and published results are examined for congruence with available ecological and distributional data. The minimum length Wagner tree and a published scheme of Polhemus show maximum congruence. A classification isomorphic with the Wagner tree is proposed as the one which should be used for future studies in the Leptopodomorpha. Objective criteria for evaluating classifications and properties of cladistic and phenetic methods are discussed. [Cladistics; Wagner; phenetics; UPGMA; classification; naturalness; prediction; stability; Leptopodomorpha.] Earlier claims by pheneticists (cf. Sokal and Sneath, 1963, and contained references) that their methods would make taxonomy logical and objective and by Hennig (1966) that the phylogenetic system of classification should serve as the general reference system for biology precipitated an extended debate about the goals and methods of biological classifi-cation. Much of the debate has not served to resolve the fundamental issue of "What is the best classification for some or all organisms and how might it be determined?" Rather, it has often concentrated on terminology without reference to concepts (cf. Mayr, 1974; Ashlock, 1979 [see Wiley, 1979]) or on motivational concerns as, for example, expressed by Bock (1977) or Ashlock (1979) who wish to make maximum use of evolutionary theory in biological classification, or Sokal and Sneath (1963) and Sneath and Sokal (1973) who prefer not to interpret classification in a phylogenetic sense (Farris, 1977). Some have wondered if we can or should arrive at a best classification (Frelin and Vuilleumier, 1979) and others have claimed simply that there is no best classification (Johnson, 1970; Key, 1974). Classifications are based on the analysis of data, which can be in the form of discrete characters or distance information. Quantitative techniques are available for analyzing both types of data (Farris, 1970, 1972; Sneath and Sokal, 1973). The interpretations we give to results are primarily related not to the way we col-

Stripes do not a zebra make, part i: a cladistic analysis of equus

Abstract: Bennett, D. K. (Department of Systematics and Ecology, The University of Kansas and Museum of Natural History, Lawrence, Kansas 66045) 1980. Stripes do not a zebra make, part I: A cladistic analysis of Equus. Syst. Zool., 29:282-287.-Living and extinct species in Equus have not been reviewed for nearly a century. More than twenty morphological, mostly cranial characters of Equus and Dinohippus are here explained; synapomorphies uniting Equus and Dinohippus and autapomorphies within Equus are discussed. Significant outgroups compared are Dinohippus Quinn, Astrohippus Stirton, and Neohipparion Gidley. Cladistic analysis indicates that Equus is a monophyletic taxon closely related to Dinohippus, and that Equus can reasonably be divided into two (and only two) subgenera, Equus (Equus) and Equus (Asinus), each of which is characterized by a suite of autapomorphic features. The North American fossil record contains close relatives of every living species of equid except E. quagga. An examination of the zoogeographic implications of the cladistic hypothesis here presented indicates a complex pattern of migration from North America to Eurasia during Blancan through late Pleistocene time, and a strong zoogeographic relationship between Africa and North America demonstrated by the equids. [Cladistics; Equus; zoogeography; fossil horses; phylogeny.] The living and extinct species of the genus Equus have not been reviewed

Evolutionary Genetics of Birds II. Conservative Protein Evolution in North American Sparrows and Relatives

Abstract: Avise,J C,J C Patton, and C F Aquadro (Departments of Zoology and Genetics, University of Georgia, Athens, Georgia 30602) 1980 Evolutionary genetics of birds II Conservative protein evolution in North American sparrows and relatives Syst Zool, 29:323-334-Differentiation at 20-21 protein-coding genes was examined by conventional techniques of starch-gel electrophoresis among twelve species and seven genera of North American sparrows and relatives, Emberizidae, subfamily Emberizinae One species representing Fringillidae was also included Data were summarized in a distance matrix which was subsequently used to infer phylogenetic trees by a variety of methods Results were generally consistent with current classification Two salient results were unanticipated: 1) the relatively close genetic similarity of Pipilo to group I Emberizinae; 2) the relatively large genetic distance of Calcarius from other Emberizinae A search of the literature revealed that the distribution of a behavioral characteristic, "bilateral scratching," had led to a prediction of phylogenetic relationships for these genera fully consistent with the protein information This result is significant because it lends support to proposals that some behavioral traits are extremely valuable phylogenetic markers Levels of protein divergence in birds are compared to previous estimates for other vertebrate taxa At corresponding levels of the taxonomic hierarchy, birds consistently exhibit far smaller genetic distances than do many fishes and other vertebrates [North American sparrows; Emberizidae; Fringillidae; protein evolution; electrophoresis; evolutionary rela-

A critique of the punctuated equilibria model and implications for the detection of speciation in the fossil record

Abstract: Levinton, J. S., and C. M. Simon (Department of Ecology and Evolution, State University of New York, Stony Brook, New York 11794) 1980. A critique of the punctuated equilibria model and implications for the detection of speciation in the fossil record. Syst. Zool., 29: 130-142.-The evolutionary models of punctuated equilibria and species selection: 1. rely upon a model of species origin from peripheral isolates, and 2. interpret trends as the net result of selection among daughter species, whose morphologies are random with respect to the trend. The punctuational model asserts that gradual morphological change is less important than the sudden and rapid morphological change which occurs at speciation, and that morphological stasis is the rule through most of the duration of a species' history. Several considerations suggest limitations of these models. 1. Peripheral population model-Evolutionary biologists acknowledge a diversity of speciation models. The extinctionof-intermediates allopatric model and the parapatric model do not require peripheral isolates, and suggest that morphological differentiation may likely arise from typical within-species geographic variation. 2. Anagenesis versus speciation-The punctuated equilibria model, by establishing a dichotomy between evolution at speciation and evolution before and after speciation, obscures more about the tempo and mode of evolution than it clarifies. 3. StasisThe nature of paleontological species level taxonomy requires the identification of speciesspecific characters which are invariant with time. This confounds the identification of stasis with species identification. Perhaps a comparison of within- versus among-species character variation might suggest whether within-species variation is the "stuff' of larger scale evolution. 4. Species selection-This requires that morphological characteristics of daughter species be random with respect to a long term trend. Ontogenetic and functional morphological interpretations of phylogeny suggest that trends within species are not necessarily random with respect to trends among species. We describe an example (scallops) where the range of possible daughter species is very restricted. We finally describe a test for the effect of average species duration on rate of anagenesis. Comparing species longevities does not adequately predict the extent of morphological divergence. [Punctuated equilibria; speciation; anagenesis; gradualism; species selection.]

Planktonic Microfossils and the Recognition of Ancestors

Abstract: erally be recognized. We can tell if absence of a form from an area is real or due to lack of preservation. Piston cores from the entire world ocean are available for all the water masses since the late Miocene. Their stratigraphy can be carefully worked out by at least three independent means of correlation, and in some cases these cores faithfully record events spaced only hundreds of years apart. Thus, in certain pelagic microfossils it is possible to sample all populations that have existed through millions of years, and be confident that no forms of interest remain unsampled. Therefore, the marine microfossil record can meet the criteria necessary to recognize ancestors. An example from the Radiolaria is discussed. [Biogeography; microfossils; ancestor/descendant; Radiolaria; stratigraphy; cladistics; phylogeny.]

Anolis lizards of the eastern caribbean: a case study in evolution. iii. a cladistic analysis of albumin immunological data, and the definition of species groups

Abstract: Gorman, George C., Donald G. Buth, and Jeff S. Wyles (Department of Biology, University of California, Los Angeles, California 90024) 1980. Anolis lizards of the eastern Caribbean: A case study in evolution. III. A cladistic analysis of albumin immunological data, and the definition of species groups. Syst. Zool., 29:143-158.-Albumin immunological distance data obtained primarily from the literature are interpreted cladistically for representatives of virtually all recognized groups of eastern Caribbean Anolis. A classification is proposed that differs substantially from the classification of Williams (1976). We argue that our classification is highly concordant with the character states used by Williams to generate his genealogy, but his interpretation cannot be forced into compatibility with the albumin evidence. We make no assumptions about regularity of rates of the albumin molecule, but the null hypothesis that rates of albumin evolution are equivalent along each lineage cannot be falsified. [Micro-complement fixation; immunological distance; cladistics; Anolis.] With well over 200 described species, the lizard genus Anolis can be very taxing to the working taxonomist. The current classification of the genus rests primarily on interpretation of data from the detailed comparative osteological study of Etheridge (1960). Williams (1976) used these data as a basis for presenting a formal classification of West Indian Anolis. In the terminology of Williams (1976), the genus is divided into two sections termed a and /8. These sections are distinguished by the absence (a) or pres