The use of chromosomal data in the systematics of viviparous onychophorans from Australia (Onychophora: Peripatopsidae)

A chromosomal analysis of populations of viviparous Australian onychophorans has uncovered a large radiation in karyotypic form. Chromosome numbers of 18, 26, 30, 32, 33 and 34 were observed, and these classes could be further subdivided on the basis of internal size relationships. Given the practical difficulties with the systematics of this group, the use of chromosomal data promises to be particularly enlightening. This is the first time any karyotypic data have been published for onychophorans since a single species was analysed in 1900.     

Onychophora in invertebrate conservation: priorities, practice and prospects

The rationale of promoting onychophorans as target taxa in invertebrate conservation is discussed. Current knowledge relevant to onychophoran conservation is outlined and gaps are identified. The prospects for conserving onychophorans are discussed, and priorities suggested.     

Genetic differentiation within New Zealand Onychophora and their relationships to the Australian fauna

Specimens of peripatopsid Onychophora from New Zealand (the viviparous Peripatoid.es novaezealandiae and P. suteri and the oviparous Ooperipatellus insignis) have been analysed by allozyme electrophoresis and compared with three species of viviparous peripatus and 0. insignis from Australia. The data indicate that at least two species of Peripatoides are present, in addition to those described. Populations collected from Dunedin, in the South Island, are not referable on morphological grounds to any described genus. Furthermore, specimens exhibit such a high level of genetic distinctiveness that we speculate they have diverged from the Peripatoides and Euperipatoides lineages prior to the separation of Australia and New Zealand some 95 Myr ago. Specimens referable to 0. insignis from New Zealand are genetically quite distinct from 0. insignis at its type locality in Australia and must be considered endemic.     

A world checklist of Onychophora (velvet worms), with notes on nomenclature and status of names

Currently, the number of valid species of Onychophora is uncertain. To facilitate taxonomic work on this understudied animal group, we present an updated checklist for the two extant onychophoran subgroups, Peripatidae and Peripatopsidae, along with an assessment of the status of each species. According to our study, 82 species of Peripatidae and 115 species of Peripatopsidae have been described thus far. However, among these 197 species, 20 are nomina dubia due to major taxonomic inconsistencies. Apart from nomina dubia, many of the valid species also require revision, in particular representatives of Paraperipatus within the Peripatopsidae, and nearly all species of Peripatidae. In addition to extant representatives, the record of unambiguous fossils includes three species with uncertain relationship to the extant taxa. For all species, we provide a list of synonyms, information on types and type localities, as well as remarks on taxonomic and nomenclatural problems and misspellings. According to recent evidence of high endemism and cryptic speciation among the Peripatidae and Peripatopsidae, previous synonyms are revised. Putative mutations, subspecies and variations are either raised to the species status or synonymised with corresponding taxa. In our revised checklist, we follow the rules and recommendations of the International Code of Zoological Nomenclature to clarify previous inconsistencies.     

Sexual dimorphism in Australian Echiniscus (Tardigrada, Echiniscidae) with descriptions of three new species

The presence of males in species of Echiniscus has been reported only as recently as 1987 and is still considered to be rare. However, males have been found to be common in a disproportionately large number of species of this genus found in Australia. Evidence of sexual dimorphism is presented for five species (three of which are new to science), females and males of which are differentiated on the basis of distinctive gonopores. In all five species males are shorter with longer clavae. Males of Echiniscus jamesi sp. nov., Echiniscus rodnae sp. nov. and Echiniscus curiosus sp. nov. have significantly longer claws. In E. rodnae sp. nov. and E. curiosus sp. nov. males have significantly longer lateral trunk appendages and the dorsal trunk appendage combinations of males of the latter are different from those of females. The widespread distribution of bisexual species of Echiniscus in xeric habitats in Australia does not support the current theory that parthenogenetic species are favoured in such environments.     

Adaptive, pre-adaptive and non-adaptive components of radiations in ancient lakes: a review

Ancient lakes are ideal model systems for evolutionary studies, as they hold hundreds of endemic species. The vast majority of these still occur in the cradle of their origin. We distinguish three different modes of speciation (allo-, para- and sympatric) which have occurred in these habitats. Although radiations from ancient lakes are generally assumed to be adaptive, we cannot fully support this point of view, because non-adaptive radiations also appear to be common, for example through chromosomal changes, hybridization or sexual selection. Even in supposedly adaptive cladogenesis, e.g. as concerns the presumed trophic adaptations of cichlid (Pisces) mouth and tooth shapes, both adaptive and non-adaptive components are acting. Distribution patterns of non-marine ostracods (Crustacea) within and outside of ancient lakes indicate that sexual reproduction might be an additional requirement for successful radiations in ancient lakes, at least in certain groups. This can best be understood by invoking ecology-based hypotheses on the evolutionary superiority of sexual reproduction such as Fisher–Muller accelerated evolution and the Tangled Bank.     

Allozyme genetic distances and evolutionary relationships in species of akodontine rodents (Cricetidae: Sigmodontinae)

Allozyme genetic distances were estimated for ten species of akodontine rodents, as compared with the Oryzomyini Oligoryzomys longicaudatus , which was used as an outgroup to assess plesiomorphic character-states. Twenty-six loci were analysed. Distribution patterns of allele frequencies were determined by both phenetic (UPGMA) and cladistic (PAUP') techniques. Allozyme analysis confirmed monophyly for the Akodontini, and among them, the distinctiveness of the genus Oxymycterus. Genetic divergence among the eight species of Akodon was small compared to most known species of rodents. Phenrtic and phylogenetic analysis between Bolonys obscurus and species of Akodon was m agreement with previous chromosomal work but in disagreement with the indications of morphology. The general lack of allozymic differentiation among members of the Akodontini suggests that in this group molecular divergence is unrelated to speciation. Molecular clock estimation calibrated by fossils showed that generic divergence within Akodontini started at least in the late Miocene and that divergence of Akodintini from Orizomyini occurred within the Miocene.     

Allozyme analysis and phyletic relationships of two new stick-insects from north-west Sicily: Bacillus grandii benazzii and B. rossius-grandii benazzii (Insecta Phasmatodea)

Two new Sicilian stick-insects have been discovered within the genus Bacillus. Evidence concerning the subspecific differentiation of Bacillus grandii benazzii and the hybrid constitution of B. rossius-grandii benazzii by means of allozyme analysis is given, and is consistent with morphological and karyological data on the differentiation of these two north-west Sicilian morphs from south-eastern B. g. grandii and B. whitei. B. g. benazzii is more polymorphic than B. g. grandii and B. rossius-g. benazzii embodies its genetic variability, thus differing sharply from the south-eastern parallel hybrid B. whitei ( = B. rossius × B. g. grandii). Reproductive mechanisms also appear to be different in the two interspecific hybrids, involving parthenogenesis in the latter and hybridogenesis in the former. Finally, the phyletic relationships among all Sicilian Bacillus taxa are summarized in a revised scheme, which also takes into account new evidence that B. atticus has contributed to hybrid constitution of the triploid B. lynceorum.     

Ultrastructure of mesoderm in embryos of Opisthopatus roseus (Onychophora, Peripatopsidae): revision of the "long germ band" hypothesis for Opisthopatus

In previous studies, an unusual pattern of development which resembles the "long germ band" development of some insects has been described in the onychophoran Opisthopatus cinctipes. This pattern has been proposed to be a characteristic of the genus Opisthopatus. To test this assumption, the ultrastructure of embryos of O. roseus, the sister species of O. cinctipes, was examined. Two kinds of paired, segmentally arranged coelomic cavities were found in the embryos studied: 1) dorsolateral coelomic cavities lined by extremely thin epithelia, and 2) ventral coelomic cavities situated within the anlagen of ventrolateral body appendages. Only the dorsolateral coelomic cavities can be considered "somites," since they occur earlier during embryogenesis. This is in contrast with the previous view that suggested a ventral position of "somites" in O. cinctipes. In addition, an anterior-to-posterior gradient occurs in the development of O. roseus. Based on our findings, we reevaluated the previous data on O. cinctipes. From this survey, no evidence in support of a "long germ band" hypothesis in Opisthopatus was found. Instead, the embryogenesis in representatives of Opisthopatus is more similar to that in other onychophorans than expected.     

Native and introduced Onychophora in Singapore

Both families of Onychophora apparently occur in Singapore. Although records of the peripatid Eoperipatus sumatranus (Sedgwick, 1888) extend over a number of years, peripatopsids were unknown there until specimens, one of which is identified as Paraperipatus papuensis (Sedgwick, 1910), were found near the Singapore Botanic Gardens. They were probably carried in the fern fibre packing of plants imported from New Guinea. Records of Onychophora in New Guinea are reviewed, and among those that have not previously been published are important collections from the 1938—1939 (3rd) Archbold Expedition to New Guinea. Other possible cases of passive transport of Onychophora are discussed.     

Protein electrophoretic variability in Saimiri and the question of its species status

An electrophoretic survey of 15 protein systems (22 loci) was employed to determine the genetic relationships among 9 populations (441 individuals) of South American squirrel monkeys (Saimiri sciureus sciureus, S. sciureus boliviensis, and S. sciureus ustus). Genetic markers capable of differentiating the second from the two other taxa were observed mainly in the ADA and GPI systems. Heterogeneity for ADA and CA2 between populations from opposite banks of the Jamari river was verified in S. sciureus ustus. The average heterozygosities ranged from 3% to 5%, the lowest being in S. sciureus sciureus and the highest in S. sciureus boliviensis. Low genetic distances (D = 0.001?0.057) were observed between populations within taxa or between S. sciureus sciureus and S. sciureus ustus. But both differed to a larger extent from S. sciureus boliviensis (D = 0.11 in both comparisons). There is a positive correlation between the genetic and geographic distance matrices. The three taxa are more clearly separated (D = 0.76–0.77) from the outgroup used for comparison (Cebus apella). Our data suggest that there is only one large, polytypic species of squirrel monkeys in South America, S. sciureus, forming a contiguous ring of geographical races or subspecies. Two of the most differentiated forms meet at the Peruvian Amazonia where natural hybrids and secondary intergradation have been reported. © 1993 Wiley-Liss, Inc.     

Islands in the sea: extreme female natal site fidelity in the Australian sea lion, Neophoca cinerea

Pinnipeds (seals, fur seals, sea lions and walrus) form large breeding aggregations with females often remaining faithful to a natal site or area. In these cases, females are philopatric to regional areas on broad geographical scales of hundreds to thousands of kilometers. An investigation of variation in a control region sequence of mtDNA in the Australian sea lion (Neophoca cinerea) has shown a case of extreme female natal site fidelity that has resulted in almost fixed population differentiation across its range (PhiST=0.93). This high level of population subdivision over short geographical distances (approx. 60 km) is unparalleled in any social marine mammal and reflects the unique life-history traits of this rare species. The high level of population subdivision and exclusive female natal site fidelity has important ramifications for conservation management, and poses many interesting questions of both academic and applied interest.     

Allozyme Variation and the Genetic Structure of Populations of the Rare Sedge Carex misera (Cyperaceae)

Abstract Carex misera is a rare sedge, endemic to rocky outcrops and mountain summits within the southern Appalachian Mountains from northern Georgia to northern North Carolina. We assessed allozyme diversity for 406 individuals from nine populations over most of the geographic range. Twenty-seven putative loci were assayed and eight (30%) were found to be polymorphic. Nei's gene diversity statistics ( H _T =0.043, H _S =0.019, G _ST =0.551) indicated low levels of variation but relatively highly differentiated populations, suggesting little gene flow. Significant deviations from genotypic expectations under Hardy-Weinberg equilibrium, high positive fixation indices, and the existence of small genetic neighborhoods within populations suggest that at least some inbreeding occurs. Cluster analysis of Nei's genetic identity statistics and principal component analysis of allele frequency data showed high similarity among the six southern populations with the two northern populations more differentiated from them and from each other. These results suggest that preservation of the northern populations is necessary to conserve the already low levels of genetic diversity within the species.     

Allozyme variation in the endangered insular endemic Castilleja grisea

BACKGROUND AND AIMS: Genetic diversity in Castilleja grisea, an endangered, perennial herb endemic to San Clemente Island, California was investigated. Subsequent to the elimination of goats from the island in 1992, many populations of C. grisea have reappeared and have been increasing in size. METHODS: Nineteen populations were surveyed for their genotype at 19 allozyme loci. KEY RESULTS: At the taxon level, 57.9 % of loci are polymorphic with A(P) = 3.09 and H(E) = 0.137. Populations averaged 33.0 % polymorphic loci with A(P) = 2.43 and H(E) = 0.099. Most variation is found within rather than among populations (G(ST) = 0.128), although differentiation among populations is significant. Genetic identities range from I = 0.960 to I = 1.000 with mean I = 0.990. There is no significant relationship between genetic and geographic distance. Gene flow among populations is Nm = 2.50 based on private alleles and Nm = 1.70 based on F(ST). Outcrossing rates based on fixation indices average t = 1.01, indicating a primarily out-crossed mating system. CONCLUSIONS: The observed genetic variation is moderately high, unusually so for an insular endemic species, suggesting that C. grisea may not have lost substantial genetic variation during 150 years of overgrazing, and indicating that it is unlikely to be endangered by genetic factors.     

Subterranean phylogeography of freshwater crayfishes shows extensive gene flow and surprisingly large population sizes

Subterranean animals are currently viewed as highly imperiled, precariously avoiding extinction in an extreme environment of darkness. This assumption is based on a hypothesis that the reduction in visual systems and morphology common in cave faunas reflects a genetic inability to adapt and persist coupled with the perception of a habitat that is limited, disconnected, and fragile. Accordingly, 95% of cave fauna in the United States are presumed endangered due to surface environmental degradation and limited geographic distributions. Our study explores the subterranean phylogeography of stygobitic crayfishes in the southeastern United States, a global hotspot of groundwater biodiversity, using extensive geographic sampling and molecular data. Despite their endangered status, our results show that subterranean crayfish species have attained moderate to high levels of genetic diversity over their evolutionary histories with large population sizes and extensive gene flow among karst systems. We then compare the subterranean population histories to those of common surface stream-dwelling crayfishes. Our results show recent drastic declines in genetic variability in the surface crayfish and suggest that these species also warrant conservation attention.     

Introduction. Speciation in plants and animals: pattern and process

Although approximately 150 years have passed since the publication of On the origin of species by means of natural selection, the definition of what species are and the ways in which species originate remain contentious issues in evolutionary biology. The biological species concept, which defines species as groups of interbreeding natural populations that are reproductively isolated from other such groups, continues to draw support. However, there is a growing realization that many animal and plant species can hybridize with their close relatives and exchange genes without losing their identity. On occasion, such hybridization can lead to the origin of new species. A key to understanding what species are and the ways in which they originate rests to a large extent on a detailed knowledge of the nature and genetics of factors that limit gene flow between species and the conditions under which such isolation originates. The collection of papers in this issue addresses these topics and deals as well with some specific issues of hybrid speciation and the causes of species radiations. The papers included arise from a 1-day symposium on speciation held during the Sixth Biennial Meeting of the Systematics Association at Edinburgh in August 2007. In this introduction, we provide some background to these papers and highlight some key points made. The papers make clear that highly significant advances to our understanding of animal and plant speciation are currently being made across the range of this topic.     

Hybridization and introgression in Carpobrotus spp. (Aizoaceae) in California. II. Allozyme evidence

Morphological evidence has indicated that hybridization and introgression are occurring between Carpobrotus edulis L., a nonindigenous, invasive species in California (Bolus), and its putative native congener, C. chilensis. The identification of allozyme markers has enabled us to quantify hybridization and the extent of introgression between these species. Samples from 20 individuals of each of five morphological types (both parent species and three hybrid phenotypes) were collected from 39 populations along the coast of California from the Mexico to Oregon borders. Ten enzyme systems revealed a total of 17 resolvable loci, eight of which were polymorphic for the genus. Five of the polymorphic loci easily differentiate C. edulis and C. chilensis. Allele frequencies among the morphologically defined types are consistent with estimations of allele frequency changes due to first- and second-generation backcrossing. In comparison to long-lived, herbaceous perennials and widespread species, C. edulis and intermediate types have more variation in their populations (P = 41.73, Ap = 2.11, Ho = 0.246, and P = 38.13, Ap = 2.06, Ho = 0.216, respectively) than C. chilensis (P = 11.76, Ap = 2.00, Ho = 0.082). Indirect estimates of gene flow indicate four of the five morphological types are outcrossing. Gene flow between previously allopatric species may have broad implications if it results in an increase in fitness; further experimentation is needed to determine the ultimate ecological consequences of this phenomenon and its possible threat to limited, remaining natural habitat in California.     

Molecular phylogeny reveals extensive ancient and ongoing radiations in a snapping shrimp species complex (Crustacea,Alpheidae, Alpheus armillatus)

Tropical marine habitats often harbor high biodiversity, including many cryptic taxa. Though the prevalence of cryptic marine taxa is well known, the evolutionary histories of these groups remain poorly understood. The snapping shrimp genus Alpheus is a good model for such investigations, as cryptic species complexes are very common, indicating widespread genetic diversification with little or no morphological change. Here, we present an extensive phylogeographic investigation of the diversified amphi-American Alpheus armillatus species complex, with geographic sampling in the Caribbean Sea, Gulf of Mexico, Florida, Brazil, and the tropical Eastern Pacific. Sequence data from two mitochondrial genes (16SrRNA and cytochrome oxidase I) and one nuclear gene (myosin heavy chain) provide strong evidence for division of the species complex into six major clades, with extensive substructure within each clade. Our total data set suggests that the A. armillatus complex includes no less than 19 putative divergent lineages, 11 in the Western Atlantic and 8 in the Eastern Pacific. Estimates of divergence times from Bayesian analyses indicate that the radiation of the species complex began ～10 MYA with the most recent divergences among subclades dating to within the last 3 MY. Furthermore, individuals from the six major clades had broadly overlapping geographic distributions, which may reflect secondary contact among previously isolated lineages, and have apparently undergone several changes in superficial coloration, which is typically the most pronounced phenotypic character distinguishing lineages. In addition, the extensive substructure within clades indicates a great deal of molecular diversification following the rise of the Isthmus of Panama. In summary, this investigation reflects substantial biodiversity concealed by morphological similarity, and suggests that both ancient and ongoing divergences have contributed to the generation of this biodiversity. It also underlines the necessity to work with the most complete data set possible, which includes comprehensive and wide-ranging sampling of taxa.     

Evidence for the presence of two sympatric species of mice (genus Mus L.) in southern France based on biochemical genetics

Populations of mice established outdoors as well as indoors have been investigated at 24 loci using starch gel electrophoresis. Two reproductively isolated groups are recognized, one of which is referable to a house mouse subspecies, Mus musculus brevirostris, and the other to a different species, Mus spretus, contrary to the view of Schwarz and Schwarz that only one species of Mus is present in the Mediterranean Basin. The genetic distance between these two groups is larger than between any pair of investigated subspecies of M. musculus. M. m. brevirostris is biochemically almost indistinguishable from M. m. domesticus. On the other hand, M. spretus exhibits several allelic variants unknown or at most very infrequent in M. musculus, as for instance at the lactate dehydrogenase B-chain locus.This work was supported by research grants from the Centre National de la Recherche Scientifique (E.R.A. No. 261) and the Ecole Pratique des Hautes Etudes.