Relict Endemics Research Articles

The Role of Old Relicts in Structuring the Boreal/Tropical Transitional Zone: The Case of East Asian Planktonic Cladocera (Crustacea)

Previous studies provided evidence of the mixing of boreal–tropical cladocerans (Crustacea) in the Far East of Eurasia, as well as the presence of numerous pre-Pleistocene relict endemics. In this study, we hypothesize that the colonization history is reflected in the proportions of endemic/boreal/tropical taxa among different habitat types within this region. We analyzed 442 qualitative samples collected from seven sub-regions of the Far East of Russia and South Korea along a latitudinal transect of approximately 2200 km, where we identified 101 Cladocera species. Our results showed a significantly higher proportion of endemic taxa in the plankton compared to the littoral and benthic zones. The proportions of endemic/boreal/tropical taxa in the permanent and temporary waters were similar. We observed a distinct shift in dominance in benthos and littoral zone across our transect: boreal taxa were prevalent in the northern sub-regions (1–4), while tropical taxa were prevalent in the southern sub-regions (5–7), with a pronounced boreal–tropical transition zone (sub-regions 4–5). Our findings suggest that this pattern arose due to the unhindered dispersion of the two faunas in a latitudinal direction until they met at the boundary during their arrival in the Late Pleistocene. In contrast, the proportions of the boreal and tropical taxa in plankton did not exceed 16% in any of the studied sub-regions, and the proportion of boreal taxa decreased from 16% to 10%, while that of the tropical taxa increased from 0% to 10% from north to south. We propose that the lower proportions and the absence of mixing of fauna in the plankton can be explained by the strong presence (sometimes dominance) of relics that occupy these water bodies before the arrival of colonizers from the north and south. Our findings highlight the significant role of biotic interactions in the formation of biogeographic boundaries, which was previously underestimated.

Taluses and Downfall Slopes as Geomorphological Refugia of Flora Relict Endemics of the Mountain Crimea and their Modern Phytoindication

In geomorphology, slopes are defined as any kind of irregularities of vertical and subhorizontal surface. A slope is an inclined part of the earth’s surface, which steepness is sufficient to determine further development of relief as a single geomorphological complex over its entire surface. There were given the being heterogeneous geomorphologic structures, slopes and their azonal landscapes and the reasons for peculiarities of the ecological environment, the link of some rare species to specific relief microforms have. The study of this phenomenon is important for geomorphology, paleogeography, phytocenology, elaboration of measures for protecting rare species (relics, endemics, relic endemics).

Genetic characterization of tertiary relict endemic Phoenix theophrasti populations in Turkey and phylogenetic relations of the species with other palm species revealed by SSR markers

Phoenix theophrasti is one of the few tree species tertiary relict endemic to the eastern Mediterranean, and it is one of the two palm species native to continental Europe. It has local populations both in Crete-Greece and in southwest Turkey. Like most of the relict endemics, it has a restricted distribution and it is at risk of extinction. The primary goal of this study was the genetic characterization of Turkish P. theophrasti populations by using SSR markers. The secondary goal was to detect the phylogenetic relations of P. theophrasti with nine different palm species which are P. dactylifera, P. reclinata, P. rupicola, P. roebelenii, P. canariensis, P. loureiri, P. acaulis, P. sylvestris from Phoenix genus and Chamaerops humilis from Chamaerops genus by the same markers. Chamaerops humilis was included in the study as the second natural palm species of the Europe. The genetic differentiation coefficient (FST) was found as 0.34, and the level of gene flow (Nm) within a generation among populations was found as 0.49. In general, excess of homozygotes relative to that expected with random mating was detected in the populations. The lowest differentiation of P. theophrasti was from P. dactylifera (FST = 0.1932), and the highest differentiation of P. theophrasti was from C. humilis (FST= 0.4261). We propose that P. theophrasti must urgently be included in the Red List of IUCN under the critically endangered (CR) category. Phylogenetic relations determined among the palm species indicated the necessity of re-evaluation of the taxonomy of palms.

South-west Sri Lanka: a floristic refugium in South Asia

In South Asia, south-west Sri Lanka alone represents the sole surviving perhumid region where Mixed Dipterocarp Forests (MDF) harbour a number of relict signature taxa providing evidence of ancient plant migration routes from Gondwana to perhumid Far Eastern Sunda land, at least partially via southern Laurasia. Palynological and phylogenetic evidence now confirm that the Malesian rain forest flora overwhelmingly immigrated from tropical Africa/Madagascar and South Asia along an ever-wet equatorial corridor sometime before the collision of the Indian Plate with the Southern Laurasian coast in middle Eocene (c. 45 Ma). This ever-wet climatic and geological history has led to the evolution of a characteristic species composition and dynamics within SW Sri Lanka’s rain forest communities, which exhibit remarkable patterns of species distribution and habitat specialization, and which parallel those of Far Eastern sister taxa. We here examine the patterns of floristic variation in each of three MDF sites using classification and ordination methods to analyse tree data collected in 62 random plots, each 0.25ha, along an altitudinal gradient. In addition, a 25 ha forest dynamics plot (FDP) was established in the Sinharaja World Heritage Site, within the CTFS-ForestGEO network that addresses issues related to community ecology and phylogenetics incorporating large-scale biogeographic patterns of major clades of both plants and animals. In the above studies, ecologically distinct floristic assemblages were revealed, i) among the three MDF sites as well as, ii) among the ridge-, slope-, and valley habitats within each forest site. In the FDP too, >80% of the 125 species, with individuals >1 cm dbh and >100 individuals per species, are significantly more associated with one or more of eight topographic habitat categories than in others. These results suggest that ecological ranges and dispersion of tree species in SW Sri Lanka is primarily mediated by soil water and nutrient levels in topographically different catenal habitats mirroring those in the Far East. Many of these species are threatened relict endemics, and their distribution patterns have important conservation implications. Their spatial distributional features provide useful criteria in site-species matching or environmental filtering in forest restoration efforts.

Coleoptera from the middle-upper Eocene European ambers: generic composition, zoogeography and climatic implications

The paper contains a review of coleopteran genera known from Baltic, Bitterfeld and Rovno amber localities. Altogether 420 genera (191 extinct and 229 extant) from 78 families are listed from these three Lagerstätten (as of 7 March 2017). The listed beetles were analyzed zoogeographically and distributional maps for 72 genera were compiled. One-quarter (56) of the genera that have survived since the Eocene have cosmopolitan ranges at present; 35 extant genera have been extripated from the Palaearctic since the Eocene. Approximately 40% of beetle genera from the middle-upper Eocene European ambers can be encountered in the wild in present-day Europe, while 5 of these genera are supposed to be European relict endemics originating in Fennosarmatia. The general similarity of the Baltic amber (s.l.) beetle assemblage to modern south Palaearctic fauna is the strongest, the Nearctic elements are more numerous in the middle-upper Eocene European ambers than the Oriental taxa. The simplified Mutual Climatic Range (MCR) method was used for palaeoclimate reconstruction based on fossil beetles. The coleopteran assemblage of Baltic amber is interpreted as indicative of warm temperate, humid, equable climate with reduced thermal seasonality [annual average temperatures range from +10–20˚C; mean of the coldest month temperatures around +10˚C; mean of the hottest month temperature around +20–24˚C; annual precipitation around 750–1500 mm]. The primary importance of high humidity for existence of the Eocene biota is pointed out.

Phylogenetic relationships, genetic divergence, historical biogeography and conservation of an endangered gecko, Goniurosaurus kuroiwae (Squamata: Eublepharidae), from the Central Ryukyus, Japan.

The Kuroiwa's eyelid gecko Goniurosaurus kuroiwae is an endangered species in a state of relict endemism in the Central Ryukyus, Japan, and is divided into five subspecies. We analyzed variations in sequence data for approximately 1900 base positions of mitochondrial 12S and 16S rRNA, and cytochrome b genes from samples representing all recognized subspecies of G. kuroiwae together with those from congeneric species in order to test the relevant previous phylogenetic hypotheses and discuss biogeographical implications in the degree and pattern of genetic divergence within G. kuroiwae. Our results, while confirming a previous molecular phylogenetic hypothesis proposed on the basis of much smaller data set, negate the relationships hypothesized on morphological grounds by explicitly supporting: 1) the primary dichotomy, with substantial genetic divergence, between G. k. splendens from the Amami Island Group and the remaining subspecies all from the Okinawa Island Group; and 2) the presence of at least six independent lineages within the latter, indicating non-monophyly for two of the subspecies, G. k. kuroiwae and G. k. orientalis, in the current taxonomic definitions. The marked genetic divergence between populations of the two island groups seems to have initiated in the middle Miocene, i.e., prior to formation of straits that have consistently been separating these two island groups since the early Pleistocene. All populations of G. kuroiwae are regarded as endangered from the viewpoint of conservation genetics.

Relict Endemism of Extant Rhineuridae (Amphisbaenia): Testing for Phylogenetic Niche Conservatism in the Fossil Record

Rhineurid amphisbaenians are represented by a rich Cenozoic fossil record in North America, but today conisist of a single living species restricted to the Florida Peninsula. Such relict endemism may be the result of phylogenetic niche conservatism (PNC), the retention of ancestral traits preventing expansion into new environments. Most tests of PNC derive ancestral niche preferences from species' extant ecologies, while ignoring valuable paleontological information. To test if PNC contributes to the restricted distribution of modern Rhineura floridana, we compare the species' current environmental preferences (temperature, precipitation and soil) to paleoenvironmental data from the rhineurid fossil record. We find no evidence of PNC in modern R. floridana, as it also occurred in Florida during drier glacial periods. Ancient rhineurids also exhibit tolerance to changing climates, having undergone a shift from subtropical-humid to semi-arid savanna conditions during the Eocene-Oligocene transition. However, rhineurids nearly disappear from North America after the middle Miocene, potentially due to the onset of prolonged freezing temperatures following the mid-Miocene Climatic Optimum. This physiological limit of environmental tolerances could be interpreted as PNC for the entire family, but also characterizes much of Amphisbaenia, emphasizing the relevance of the temporal as well as phylogenetic scale at which PNC is investigated.

Relationships among the Spruces (Picea, Pinaceae) of Southwestern North America

Abstract Numerous populations from six spruce taxa, including four relict endemics, Picea chihuahuana (Chihuahua spruce), P. martinezii (Martinez spruce), P. mexicana (Mexican spruce), and P. breweriana (Brewer spruce), and two widespread species, P. engelmannii (Engelmann spruce) and P. pungens (blue spruce), were compared at homologous isozyme loci to test various hypotheses about their affinities and origins. Each of the species was clearly separated, and Neighbor-Joining and Unweighted Pair Group analyses of Nei's genetic distance grouped all populations within a taxon into their own clusters. Spruces from Flys Peak, Chiricahua Mountains, Arizona, joined a P. engelmannii cluster and were not a bridge to P. mexicana as previously believed. Spruces from Cerro Mohinora, Chihuahua, were clearly P. mexicana, not phantom hybrids of P. chihuahuana and P. pungens. Nuclear random amplified polymorphic DNA and chloroplast simple sequence repeat and cleaved amplified polymorphic genetic markers were compared in ...

Genetic variation of Saxifraga paniculata Mill. (Saxifragaceae): molecular evidence for glacial relict endemism in central Europe

Saxifraga paniculata Mill. is an amphi-atlantic species that is widely distributed in the Alps but also occurs with disjunct populations in central Europe. These isolated populations are considered to be glacial relicts. In the study presented here, we analysed the genetic variation within and between populations of Saxifraga paniculata in central Europe to test whether the molecular data give evidence for the supposed relict endemism. We used RAPD analysis and the analysis of molecular variance (AMOVA) technique to investigate 30 populations of Saxifraga paniculata from different locations in central Europe and adjacent geographical areas. In the RAPD analysis, 319 fragments could be amplified of which 91.2% were polymorphic. The percentage of polymorphic bands within populations correlated significantly with population size, indicating a higher level of inbreeding in small populations. AMOVA revealed 43.60% of the variation within populations, only 15.45% between populations within locations, and 40.95% between locations. Genetic distance between locations (ΦST) correlated significantly with the geographical distance between locations, giving evidence for geographical isolation. In an UPGMA cluster analysis, based on RAPD data, the populations were grouped together according to their location. The results exhibit a strong genetic differentiation which is obviously due to genetic drift in the isolated populations. Our study therefore gives evidence for glacial relict endemism of Saxifraga paniculata in central Europe. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80, 11–21.

Genetic Variation of Sesleria albicans Kit. ex Schultes (Poaceae): Lack of Evidence for Glacial Relict Endemism in Central Europe

Abstract: The populations of Sesleria albicans Kit. ex Schultes in central Europe are supposed to be glacial relicts. The genetic structure of these populations can provide evidence for the reliability of this hypothesis, since a distinct geographical clustering of populations and high levels of interpopulational genetic variation can be expected in the case of relict endemism. In the study presented here, we used Random Amplified Polymorphic DNA (RAPD) analysis to describe the genetic structure of 25 Sesleria albicans populations from west and central Germany, southwest Germany and the Alps. In the RAPD analysis 344 fragments could be amplified, of which 95.9 % were polymorphic. The percentage of polymorphic bands per population ranged from 29.7 % to 56.7 % and correlated significantly with population size, indicating a higher level of inbreeding in small and isolated populations. An analysis of molecular variance (AMOVA) revealed that 61.22 % of the variation resided within populations, whereas 34.25 % was found among populations within regions and only 4.53 % among regions. Genetic distance among populations (φST) varied between 0.63 and 0.13, and in a Neighbour‐Joining dendrogram based on the genetic distances between the investigated individuals, all populations could be discriminated from each other. The populations from the northern and southern Alps were clearly separated from the central European populations. These, however, did not form geographic clusters. Considering these results, Sesleria albicans showed a high level of intra‐ but only a low level of interpopulational genetic variation and very weak regional differentiation. The genetic pattern detected in this study, therefore, gives no evidence for glacial relict endemism of Sesleria albicans in central Europe.

Description of Muntiacus truongsonensis, a new species of muntjac (Artiodactyla: Muntiacidae) from Central Vietnam, and implications for conservation

AbstractA small blackish muntjac has been discovered in the west Quang Nam province of Vietnam. DNA sequencing of six individuals showed that this is a distinct species. A description is given. The new find is the third new ungulate species to be discovered in Vietnam in five years. The paper predicts that more species remain to be found in the Annamite mountain range and includes DNA analysis from two more undescribed species. The DNA analysis indicates that there has been a radiation of several related muntjac species in the Annamite mountains, also that the new truongson muntjac shows high intraspecific genetic diversity indicative of a large effective population size. The paper discusses the significance of these finds from an evolutionary and conservation viewpoint. The authors urge conservation authorities in the IndoChinese countries to devote special measures to protect this site of dynamic evolution which retains a combination of relict endemics (e.g. Pseudoryx) as well as on‐going speciation.

Two New Asian Xanthoria Species (Teloschistaceae, Lichenized Ascomycotina)

Two corticolous species of the lichen genus Xanthoria, X. alfredi S. Kondratyuk & Poelt from Russian and Chinese Far East and X. aphrodites Kalb, Poelt & S. Kondratyuk from Cyprus are described. Morphological characters, in particular the presence of true rhizines within Xanthoria and their diVerences from hapters are indicated. Xanthoria novozelandica Hillmann is the first species of Xanthoria in which oil paraphyses have been found. A key to the treated species, including X. novozelandica and Xanthoria sp. from western North America is provided. The ecology and distribution of described species are discussed. Xanthoria alfredi and X. ulophyllodes probably constitute a typical species pair; X. alfredi and X. aphrodites are relict endemics. ? 1997 The British Lichen Society

TWO NEW ASIANXANTHORIASPECIES (TELOSCHISTACEAE, LICHENIZED ASCOMYCOTINA)

Two corticolous species of the lichen genusXanthoria,X. alfrediS. Kondratyuk &Poelt from Russian and Chinese Far East andX. aphroditesKalb, Poelt & S. Kondratyuk from Cyprus are described. Morphological characters, in particular the presence of true rhizines withinXanthoriaand their differences from hapters are indicated.Xanthoria novozelandicaHillmann is the first species ofXanthoriain which oil paraphyses have been found. A key to the treated species, includingX. novozelandicaandXanthoriasp. from western North America is provided. The ecology and distribution of described species are discussed.Xanthoria alfrediandX. ulophyllodesprobably constitute a typical species pair;X. alfrediandX. aphroditesare relict endemics.

Islands: stability, diversity, conservation

Islands present both a diversity and a stability ‘paradox’. They are often highly species-poor but have considerable biological interest in terms of extraordinary endemic genera and taxonomically isolated groups. They appear to be stable, as in some cases these organisms have persisted for many millions of years, and having an oceanic climate, extreme climatic events may be comparatively rare. However, when subject to extrinsic (anthropogenic) disturbance they do not appear to be stable, but often suffer catastrophic ecological change. These apparent paradoxes are resolved when it is realized that all these features are consequences of the same island characteristics: biotic isolation and oceanicity. As a result of these two characteristics, far oceanic islands are quantitatively different from continental systems in the nature of their ecological processes, which appear to give rise to an extreme punctuated equilibrium model of evolutionary change. Endemics may be ancient relict endemics displaying prolonged stasis and persistence, or products of adaptive radiation representing rapid punctuational events. A process-based definition of a relict endemic (palaeoendemic) is one whose founding lineage (i.e. the original continental source taxon) has not left any descendents. A corollary of this definition is that the time of divergence between an endemic and its continental sister-group should predate the colonization of the island by the now endemic lineage. An example is Dicksonia arborescens which has been on St Helena for at least 9 Myrs and no longer occurs in the likely source area of Africa. These relict endemics, frequent on islands, are important as the last remnants of tranches of biodiversity that have vanished elsewhere. Island conservation strategies require an integrated understanding of both sides of the diversity and stability paradox so that both island processes and island organisms can be conserved.

The California Islands

The California Islands off the Pacific coast of southern and Baja California display almost diagrammatically the insular phenomena that characterize fringing archipelagos. To a lesser extent they also illustrate some of the insular features associated with oceanic islands. Several of these islands have been much studied and are relatively accessible. The location, size, elevation, and geological history of the islands are described. To determine the degree of disharmony of the island floras, they are analyzed in comparison with selected California mainland floras. The degree of richness of the island floras is discussed in relation to size of the islands, proximity to the mainland, and other variables. Such phenomena of island life as the immigrant pattern of vertebrate distribution, relict versus autochthonous endemism of plants and animals, insular gigantism, continuous flowering, high degree of hybridization, vulnerability, and loss of dispersibility in island species are considered in reference to the California Islands. Some suggestions as to available facilities on the islands and persons or institutions to contact are given in an epilogue. Off the Pacific coast of southern California and Baja California, Mexico, lie the islands variously called the Channel Islands or California Offshore Islands and the Baja California Islands or, collectively, the California Islands. Although they are in no sense tropical, they are worthy of mention in this symposium because they are a good example of a fringing archipelago and because they illustrate many of the insular phenomena that one might otherwise have to travel far to tropical areas to study. Some of them are quite accessible from the heavily populated areas of southern California, and some of them have been rather thoroughly studied by biologists. Although they are, thus, rather well known to California biologists, they are surprisingly little known to non-Californians. It is hoped that this short discussion of them and of some of their insular characteristics will create more interest in them among those biologists concerned with evolutionary and environmental phenomena. The Southern California and Baja California Islands consist of 16 major islands or island groups spread over some 500 miles between Point Conception, California, and Punta Eugenia, Baja California, Mexico (Fig. 1). From east to west the Northern Channel Islands are Anacapa, Santa Cruz, Santa Rosa, and San Miguel. The Southern Channel Islands are San Nicolas, Santa Barbara, Santa Catalina, and San Clemente Islands. The Baja California Islands include Los Coronados just south of San Diego, Todos Santos off Ensenada, San Martin off San Quintin, San Geronimo south of El Rosario, the San Benitos, Cedros, and Natividad off Punta Eugenia, and Guadalupe 157 miles off the Baja California peninsula. In order of size, with area in square miles, the islands are: Cedros (134), Guadalupe (98), Santa Cruz (96), Santa Rosa (84), Santa Catalina (75), San 11 am grateful to Dr. Reid Moran for critically reviewing the manuscript, for making many valuable suggestions, and for supplying useful unpublished information. 2Rancho Santa Ana Botanic Garden, Claremont, California 91711. ANN. MISSOURI BOT. GARD. 56: 391-408. 1969. This content downloaded from 157.55.39.247 on Wed, 27 Apr 2016 05:48:03 UTC All use subject to http://about.jstor.org/terms [VOL. 5 6 392 ANNALS OF THE MISSOURI BOTANICAL GARDEN Clemente (56), San Nicolas (22), San Miguel (14), Natividad (2.8), San Benito (2.5), Anacapa (1.1), Santa Barbara (1.0), Los Coronados (1.0), San Martin (0.9), Todos Santos (0.5), and San Geronimo (0.2). Their distances from the mainland, in statute miles, are: San Martin (3), Todos Santos (4), Natividad (5), San Geronimo (6), Los Coronados (8), Anacapa (13), Cedros (14), Santa Cruz (19), Santa Catalina (20), San Miguel (26), Santa Rosa (27), Santa Barbara (38), San Benito (41), San Clemente (49), San Nicolas (61), and Guadalupe (157). All these figures are from a summary by Philbrick (1967). The highest elevations on the largest islands, in feet, are: Guadalupe (4,257), Cedros (3,950), Santa Cruz (2,165), Santa Catalina (2,069), San Clemente (1,965), Santa Rosa (1,574), and San Nicolas (907).

THE SYSTEMATICS OF PHACELIA MACULATA AND P. DUBIA VAR. GEORGIANA, BOTH ENDEMIC TO GRANITE OUTCROP COMMUNITIES

Phacelia maculata and P. dubia var. georgiana are endemic to granite outcrop communities in the southeastern Piedmont of Alabama, Georgia, and South Carolina. Data are presented in support of the hypothesis that P. maculata is a relict species, whereas P. dubia var. georgiana is an ecotype recently derived from the Appalachian P. dubia. P. maculata is restricted to undisturbed outcrops of the central and upper Piedmont, and its center of distribution coincides with that of other well‐known, relict endemics. It lacks intraspecific variability and has a narrowly circumscribed habitat which has recently undergone contraction. P. dubia var. georgiana is absent from most of the undisturbed outcrops, but it occurs in abundance at outcrop sites of the lower Piedmont of Alabama and Georgia which have been recently disturbed. It is variable and will successfully compete with well‐established outcrop species in a variety of open habitats. Both its habitat and range are expanding with the increase of man's disturbance of the outcrop habitat, especially in the form of pasturing and cultivation. Each taxon is morphologically distinctive throughout its entire range, and population studies reveal no evidence of introgression in regions where they are sympatric. Reproductive isolation is complete and is effected by means of a postfertilization, incompatibility factor.