Historical biogeography at the crossroads of the northern continents: molecular phylogenetics of red-backed voles (Rodentia: Arvicolinae)

Evolutionary relationships of red-backed voles and their relatives were examined and used to test biogeographic hypotheses. Sequences of the mitochondrial cytochrome b gene were obtained for 25 individuals representing Alticola macrotis, Clethrionomys californicus, C. gapperi, C. glareolus, C. rutilus, and C. rufocanus. These were combined with 21 partial sequences from GenBank for C. regulus, C. rex, C. rufocanus, C. rutilus, Eothenomys imaizumii, E. melanogaster, Phaulomys andersoni, and P. smithii. Complete sequences of three species of Microtus (M. montanus, M. oeconomus, and M. pennsylvanicus), representative species of other arvicoline genera (Myopus, Synaptomys, Arvicola, Ellobius, Ondatra, Lemmus, Dicrostonyx, and Phenacomys), and a sigmodontine representative (Peromyscus) were included as outgroups. We used maximum parsimony, maximum likelihood, distance, and Bayesian based methods and conducted statistical tests on proposed hypotheses of phylogenetic relationships and biogeographic histories. A close relationship of species representing the genera Alticola, Clethrionomys, and Eothenomys was supported (Clethrionomyini); however, the genus Clethrionomys was paraphyletic with respect to both Alticola and Eothenomys. Three major clades were identified as Asian (Eothenomys andersoni, E. smithii, C. rex, C. regulus, and C. rufocanus), Trans-beringian (Alticola macrotis, C. californicus, C. gapperi, C. glarelolus, and C. rutilus), and Taiwanese (E. melanogaster). These results are consistent with the fossil record which indicates an initial diversification in Asia followed by colonization of the Nearctic on at least two occasions. The holarctic species, C. rutilus, appears to have either reinvaded Asia from North America or colonized North America more recently (late Pleistocene) than the two species of Clethrionomys (C. gapperi and C. californicus) that are endemic to North America (early to mid-Pleistocene). Finally, C. gapperi, appears to be comprised of an eastern and a western species, the former with affinities to the Asian C. glareolus and the latter more closely related to C. californicus.     

LisBeth: New cladistics for phylogenetics and biogeography

Within phylogenetics, two methods are known to implement cladistics: parsimony or maximum parsimony (MP) and three-item analysis (3ia). Despite the lack of suitable software, 3ia is occasionally used in systematic, and more regularly, in historical biogeography. Here, we present LisBeth, the first and only phylogenetic/biogeographic program freely available that uses the 3ia approach and offer some insights into its theoretical propositions. LisBeth does not rely on the conventional taxon/character matrix. Instead, characters are represented as rooted trees. LisBeth performs 3ia analyses based on maximum congruence of three-item statements and calculates the intersection tree (which differs from usual consensus). In biogeography, it applies the transparent method to handle widespread taxa and implements paralogy-free subtree analysis to remove redundant distributions. For the sake of interoperability, LisBeth may import/export characters from/to matrix in NEXUS format, allowing comparison with other cladistic programs. LisBeth also imports phylogenetic characters from Xper² knowledge bases.     

When phylogenetics met biogeography: Willi Hennig,Lars Brundin and the roots of phylogenetic and cladistic biogeography

Willi Hennig's (Beitr. Ent. 1960, 10, 15) Die Dipteren-Fauna von Neuseeland als systematisches und tiergeographisches Problem applied a phylogenetic approach to examine the distributional patterns exhibited by the Diptera of New Zealand. Hennig showed how phylogenetic trees may be used to infer dispersal, based on the progression and deviation rules, and also discussed the existence of vicariance patterns. The most important author who applied Hennig's phylogenetic biogeography was Lars Brundin, when analysing the phylogenetic relationships of two taxa of Chironomidae (Diptera) and using them to examine the biogeographic relationships of Australia, New Zealand, South America and South Africa. The relevance of Brundin's contribution was noted by several authors, as it began the cladistic or vicariance approach to biogeography, that implies the discovery of vicariance events shared by different monophyletic groups. Both phylogenetic and cladistic biogeography have a place in contemporary biogeography, the former for analysing taxon biogeography and the latter when addressing Earth or biota biogeography. The recent use of the term “phylogenetic biogeography” to refer to a posteriori methods of cladistic biogeography is erroneous and should be avoided.     

Linguistic diversity, global paradigms and taken-for-grantedness

In the first part of this paper taken-for-granted hypotheses in linguistic diversity are presented. In the second part the two constellations of globalized ideologies are described constituting paradigms that these hypotheses illustrate: competition and solidarity. In the third part, a condensed version of emerging globalized concerns is given. Sociolinguistics and anthropological linguistics are disciplines that increasingly theorize and analyze within the solidarity paradigm. It is suggested that a systematic uprooting of competition as taken-for-granted grounding of scientific research should allow for the development of theorization and successful applications of solidarity ideologies. In short, in this paper, our multifaceted taken-for-grantedness is challenged in many ways: (1) competition is ideological and many social movements are unmasking it by articulating solidarity as a basis for ideologies, (2) difference is not necessarily divisive but it is so in pervasive competition, (3) proponents of the Nation-State as a model of social organization have vested interests in competition, (4) competition has not favored the articulation of common human grounds but globalization helps to raise concerns and articulate commonness/solidarity in difference.     

Evolution of terrestrial birds in three continents: biogeography and parallel radiations

Aim To reconstruct the biogeographical history of a large clade of mainly terrestrially adapted birds (coraciiform and piciform birds, owls, diurnal raptors, New World vultures, trogons, mousebirds, cuckoo‐rollers, seriemas, parrots and passerines) to test the hypothesis of its Gondwanan origin. Location Global. Methods The phylogenetic tree used in the analysis was a family‐level tree estimated from previously published nuclear DNA sequence data. Each family for which a thorough and taxonomically well‐sampled phylogenetic analysis exists was subject to an initial dispersal–vicariance analysis in order to reconstruct ancestral areas for its two most basal lineages. Both basal lineages were then used to represent the family in the subsequent reconstruction of ancestral distributions for the entire radiation. Results The analysis showed that three reciprocally monophyletic groups of terrestrial birds have diversified in the Gondwanan land areas of Australia, South America and Africa, respectively. Although each of these three groups may also have originally included other groups, the only survivors today from the Australian radiation are the passerines and parrots, while the falcons and seriemas have survived from the South American radiation. The group of survivors from the African radiation is considerably more taxonomically diverse and includes all coraciiform and piciform birds, owls, diurnal raptors (except falcons), New World vultures, trogons, mousebirds and cuckoo‐rollers. Main conclusions The outlined evolutionary scenario with three geographically isolated clades of terrestrial birds is consistent with the available estimates of Late Cretaceous to early Palaeogene dates for these radiations. The diversifications and ecological adaptations within each of the three groups most likely took place in isolation on the different continents. Many cases of convergently evolved adaptations may be revealed through the increased understanding of the phylogenetic relationships of terrestrial birds.     

The influence of species diversity on ecosystem productivity: how,where, and why?

The effect of species diversity on ecosystem productivity is controversial, in large part because field experiments investigating this relationship have been fraught with difficulties. Unfortunately, there are few guidelines to aid researchers who must overcome these difficulties and determine whether global species losses seriously threaten the ecological and economic bases of terrestrial ecosystems. In response, I offer a set of hypotheses that describe how diversity might influence productivity in plant communities based on three well-known mechanisms: complementarity, facilitation, and the sampling effect. Emphasis on these mechanisms reveals the sensitivity of any diversity-productivity relationship to ecological context (i.e., where this relationship should be found); ecological context includes characteristics of the surrounding environment, temporal and spatial scales of observation, and the intensity of human management. In particular, the legitimacy of the sampling effect as a mechanism of productivity enhancement is dependent upon the degree to which stochastic events influence immigration and extinction processes in a given ecosystem. A mechanistic approach also requires that the three mechanisms be separated and quantified in diversity experiments, and I examine the most appropriate analyses for doing so, focusing on the overyielding technique. Finally, I question why productivity per se is a relevant management concern in non-agricultural systems once relationships among diversity, productivity, and the qualities of the surrounding environment are considered.     

DNA barcoding: how it complements taxonomy, molecular phylogenetics and population genetics

DNA barcoding aims to provide an efficient method for species-level identifications and, as such, will contribute powerfully to taxonomic and biodiversity research. As the number of DNA barcode sequences accumulates, however, these data will also provide a unique 'horizontal' genomics perspective with broad implications. For example, here we compare the goals and methods of DNA barcoding with those of molecular phylogenetics and population genetics, and suggest that DNA barcoding can complement current research in these areas by providing background information that will be helpful in the selection of taxa for further analyses.     

Out of Africa: molecular phylogenetics and biogeography of Wolffiella (Lemnaceae)

The monophyletic genus Wolffiella (Lemnaceae) comprises 10 species divided taxonomically into three sections. Relative to other genera of Lemnaceae, Wolffiella has a restricted range, with species distributed in warm temperate to tropical areas of Africa and the Americas, with only one species occurring in both areas. Sequence data from coding (rbcL and matK) and non‐coding (trnK and rpl16 introns) regions of cpDNA were analyzed phylogenetically to resolve relationships within Wolffiella, and these results were compared to earlier allozyme and morphological studies. Allozymes, cpDNA and morphology all supported the recognition of three sections. Relationships among species were similar in most respects between the allozyme and cpDNA trees, as well as among the different plastid partitions. In Wolffiella, both non‐synonymous and synonymous substitutions were greater in matK than in rbcL, as observed in other taxa. The synonymous substitution rate in matK was similar to the substitution rate of the non‐coding regions. All partitions, including coding regions, exhibited some homoplasy. Biogeographical reconstructions from a combination of cpDNA partitions indicated that Wolffiella originated in Africa with early movement to and radiation in the Americas. The one species found in both Africa and the Americas, W. welwitschii, likely originated in the Americas and subsequently dispersed to Africa. Using the SOWH test, the cpDNA data could reject two alternative biogeographical hypotheses suggested from analyses of morphological and allozyme data. The present distribution of Wolffiella can be explained by two major dispersal events and this contrasts with the more complex species distributions in other Lemnaceae genera. Limited dispersal in Wolffiella relative to other Lemnaceae genera may be due to more recent origins of species, lower dispersibility and poorer colonizing ability. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 79 , 565–576.     

Molecular phylogenetics, historical biogeography, and chromosome number evolution of Portulaca (Portulacaceae)

Portulaca is the only genus in Portulacaceae and has ca. 100 species distributed worldwide, mainly in the tropics and subtropics. Molecular data place the genus as one of the closest relatives of Cactaceae, but phylogenetic relationships within Portulaca are barely known. This study samples 59 species of Portulaca, 10 infraspecific taxa, and three cultivars, including multiple samples of widespread species. The sampled taxa represent all subgenera in the classifications of von Poellnitz (1934), Legrand (1958), and Geesink (1969) and come from around the world. Nuclear ITS and chloroplast ndhF, trnT-psbD intergenic spacer, and ndhA intron DNA sequences were analyzed using maximum likelihood and Bayesian methods to produce a hypothesis of relationships within Portulaca. Divergence times were estimated using Hawaiian endemics for calibration, and biogeographical patterns were examined using a Bayes-DIVA approach. In addition, the evolution of chromosome numbers in the genus was investigated using probabilistic models. The analyses strongly support the monophyly of Portulaca, with an age of the most recent common ancestor (MRCA) of 23 Myr. Within Portulaca are two major lineages: the OL clade (comprising opposite-leaved species) distributed in Africa, Asia, and Australia, and the AL clade (comprising alternate to subopposite-leaved species), which is more widespread and originated in the New World. Sedopsis, a genus sometimes recognized as distinct from Portulaca based on a long corolla tube, is nested within the OL clade and does not merit taxonomic recognition. Samples of Portulaca grandiflora, Portulaca halimoides, and Portulaca oleracea were found to be non-monophyletic. It is hypothesized that the ancestral distribution area of Portulaca included southern hemisphere continents and Asia. The OL clade remained restricted to the Old World (except Portulaca quadrifida, a pantropical weed), while the AL clade, with a South American origin, was able to disperse multiple times to other continents. The base chromosome number for Portulaca is inferred to be x=9, although the analysis was primarily based on the available data for the AL clade. A number of chromosome number change events (polyploidization, demi-polyploidization, gain, and loss) were shown to have occurred in the genus, especially within the Oleracea clade.     

Phylogeny and biogeography of the staghorn fern genus Platycerium (Polypodiaceae, Polypodiidae)

The genus Platycerium is one of the few pantropical epiphytic fern genera with six species in Afro-Madagascar, 8-11 Australasian species, and a single species in tropical South America. Nucleotide sequences of four chloroplast DNA markers are employed to reconstruct the phylogeny of these ferns and to explore their historical biogeography. The data set was designed to resolve conflicting hypotheses on the relationships within the genus that were based on previous phylogenetic studies exploring morphological evidence. Our results suggest a basal split of Platycerium into two well-supported clades. One clade comprises species occurring in Africa, Madagascar, and South America, whereas the second clade contains exclusively Australasian species. The latter clade is further divided into a clade corresponding to P. bifurcatum and its putative segregates and a clade of seven species occurring from Indochina throughout the Malesian region to New Guinea and Australia. The Afro-Madagascan clade includes a clade of two species found in tropical Africa and a clade of four species that includes three species endemic to Madagascar. The single neotropical species of this genus, P. andinum, is nested within the Afro-Madagascan clade but is not closely related to any extant species.     

Evolution and biogeography of primates: a new model based on molecular phylogenetics,vicariance and plate tectonics

Heads, M. Evolution and biogeography of primates: a new model based on molecular phylogenetics, vicariance and plate tectonics. —Zoologica Scripta, 39, 107–127. The ages of the oldest fossils suggest an origin for primates in the Paleocene (～56 Ma). Fossil‐calibrated molecular clock dates give Cretaceous dates (～80–116 Ma). Both these estimates are minimum dates although they are often ‘transmogrified’ and treated as maximum or absolute dates. Oldest fossils can underestimate ages by tens of millions of years and instead of calibrating the time‐course of evolution with a scanty fossil record, the geographical boundaries of the main molecular clades of primates are calibrated here with radiometrically dated tectonic events. This indicates that primates originated when a globally widespread ancestor (early Archonta) differentiated into a northern group (Plesiadapiformes, extinct), a southern group (Primates), and two south‐east Asian groups (Dermoptera and Scandentia). The division occurred with the breakup of Pangea in the Early Jurassic and the opening of the central Atlantic (～185 Ma). Within primates, the strepsirrhines and haplorhines diverged with volcanism and buckling on the Lebombo Monocline, a volcanic rifted margin in south‐east Africa (Early Jurassic, ～180 Ma). Within strepsirrhines, lorises and galagos (Africa and Asia) and lemurs (Madagascar) diverged with the formation of the Mozambique Channel (Middle Jurassic, ～160 Ma). Within haplorhines, Old World monkeys and New World monkeys diverged with the opening of the Atlantic (Early Cretaceous, ～130 Ma). The main aspects of primate distribution are interpreted as the result of plate tectonics, phylogeny and vicariance, with some subsequent range expansion leading to secondary overlap. Long‐distance, trans‐oceanic dispersal events are not necessary. The primate ancestral complex was already widespread globally when sea‐floor spreading, strike‐slip rifting and orogeny fractured and deformed distributions through the Jurassic and Cretaceous, leading to the origin of the modern clades. The model suggests that the topology of the phylogenetic tree reflects a sequence of differentiation in a widespread ancestor rather than a series of dispersal events.     

Evolutionary history of the parrotfishes: biogeography,ecomorphology, and comparative diversity

The family Scaridae comprises about 90 species of herbivorous coral reef, rock reef, and seagrass fishes. Parrotfishes are important agents of marine bioerosion who rework the substrate with their beaklike oral jaws. Many scarid populations are characterized by complex social systems including highly differentiated sexual stages, territoriality, and the defense of harems. Here, we test a hypothesis of relationships among parrotfish genera derived from nearly 2 kb of nuclear and mitochondrial DNA sequence. The DNA tree is different than a phylogeny based on comparative morphology and leads to important reinterpretations of scarid evolution. The molecular data suggest a split among seagrass and coral reef associated genera with nearly 80% of all species in the coral reef clade. Our phylogenetic results imply an East Tethyan origin of the family and the recurrent evolution of excavating and scraping feeding modes. It is likely that ecomorphological differences played a significant role in the initial divergence of major scarid lineages, but that variation in color and breeding behavior has triggered subsequent diversification. We present a two-phase model of parrotfish evolution to explain patterns of comparative diversity. Finally, we discuss the application of this model to other adaptively radiating clades.     

Evolution and the latitudinal diversity gradient: speciation, extinction and biogeography

A latitudinal gradient in biodiversity has existed since before the time of the dinosaurs, yet how and why this gradient arose remains unresolved. Here we review two major hypotheses for the origin of the latitudinal diversity gradient. The time and area hypothesis holds that tropical climates are older and historically larger, allowing more opportunity for diversification. This hypothesis is supported by observations that temperate taxa are often younger than, and nested within, tropical taxa, and that diversity is positively correlated with the age and area of geographical regions. The diversification rate hypothesis holds that tropical regions diversify faster due to higher rates of speciation (caused by increased opportunities for the evolution of reproductive isolation, or faster molecular evolution, or the increased importance of biotic interactions), or due to lower extinction rates. There is phylogenetic evidence for higher rates of diversification in tropical clades, and palaeontological data demonstrate higher rates of origination for tropical taxa, but mixed evidence for latitudinal differences in extinction rates. Studies of latitudinal variation in incipient speciation also suggest faster speciation in the tropics. Distinguishing the roles of history, speciation and extinction in the origin of the latitudinal gradient represents a major challenge to future research.     

A world-wide perspective on crucifer speciation and evolution: phylogenetics,biogeography and trait evolution in tribe Arabideae

Background and Aims

Tribe Arabideae are the most species-rich monophyletic lineage in Brassicaceae. More than 500 species are distributed in the majority of mountain and alpine regions worldwide. This study provides the first comprehensive phylogenetic analysis for the species assemblage and tests for association of trait and characters, providing the first explanations for the enormous species radiation since the mid Miocene.

Methods

Phylogenetic analyses of DNA sequence variation of nuclear encoded loci and plastid DNA are used to unravel a reliable phylogenetic tree. Trait and ancestral area reconstructions were performed and lineage-specific diversification rates were calculated to explain various radiations in the last 15 Myr in space and time.

Key Results

A well-resolved phylogenetic tree demonstrates the paraphyly of the genus Arabis and a new systematic concept is established. Initially, multiple radiations involved a split between lowland annuals and mountain/alpine perennial sister species. Subsequently, increased speciation rates occur in the perennial lineages. The centre of origin of tribe Arabideae is most likely the Irano-Turanian region from which the various clades colonized the temperate mountain and alpine regions of the world.

Conclusions

Mid Miocene early diversification started with increased speciation rates due to the emergence of various annual lineages. Subsequent radiations were mostly driven by diversification within perennial species during the Pliocene, but increased speciation rates also occurred during that epoch. Taxonomic concepts in Arabis are still in need of a major taxonomic revision to define monophyletic groups.     

The historical biogeography of sturgeons (Osteichthyes: Acipenseridae): a synthesis of phylogenetics, palaeontology and palaeogeography

Abstract. The historical biogeography of sturgeons is explored using information from palaeogeography, palaeontology and phylogenetic interrelationships. The integration of information from these diverse sources indicates that sturgeons reached a wide Laurasian distribution in the Cretaceous and Tertiary by freshwater and coastal dispersal routes across land connections and along newly forming continental margins. The fossil record also suggests a considerable degree of morphological stasis and also supports an estuarine habit, and perhaps diadromy, as an old and conserved life history trait. While a ‘centre of origin’ for sturgeons remains elusive, phylogenetic relationships indicate that diversification appears to have been associated with fragmentation of biota, and of landmasses and basins, by late Tertiary geological and climatic phenomena, such as orogeny and unequal glaciation over North America, the desiccation of central Asia and alteration of its drainages, and the formation of discrete Ponto-Caspian basins by the fragmentation of the Paratethys. Amphi-oceanic distributions of certain species (Acipenser medirostris Ayres) and sister taxa (e.g. A. oxyrhynchus Mitchill and A. sturio L.) are explained by coastal dispersal and subsequent vicariance by geological (sea-floor spreading and development of new continental margins) and climatic (Pliocene cooling) changes during the Tertiary. An hypothesis is developed for the relationships of the North American sturgeons and their potential relationships with the Siberian sturgeon A. baeri. Late Tertiary climatic and geological phenomena are hypothesized as mediators of vicariance and subsequent diversification of these acipenserids. It appears that although acipenserids are a geologically old group, the historical biogeography of surviving lineages is best explained by more recent geological and climatic changes.     

Invaders or endemics? Molecular phylogenetics,biogeography and systematics of Dreissena in the Balkans

1. Zebra mussels and their relatives (Dreissena spp.) have been well studied in eastern, central and western Europe as well as in North America, because of their invasiveness and economic importance. Much less is known about the biology and biogeography of indigenous (endemic) taxa of Dreissena, in the Balkans. A better knowledge of these taxa could help us (i) understand the factors triggering invasiveness in some taxa and (ii) identify other potentially invasive species. 2. Using a phylogenetic approach (2108 base pairs from three gene fragments), Dreissena spp. from natural lakes in the Balkans were studied to test whether invasive Dreissena populations occur in such lakes on the Balkan Peninsula, whether Dreissena stankovici really is endemic to the ancient Lakes Ohrid and Prespa, and to infer the phylogenetic and biogeographical relationships of Balkan dreissenids. 3. No invasive species of Dreissena, such as Dreissena polymorpha, were recorded. The supposedly ‘endemic’D. stankovici is not restricted to the ancient Lakes Ohrid and Prespa, but is the most widespread and dominant species in the west‐central Balkans. Its southern sister taxon, Dreissena blanci, occurs sympatrically with D. stankovici in Lakes Prespa, Mikri Prespa and Pamvotis. Both species are classified into the subgenus Dreissena (Carinodreissena) of which the subgenus Dreissena (Dreissena) (which includes the invasive D. polymorpha) is the sister taxon. Dreissena blanci and D. stankovici are considered to represent distinct species. 4. On a global scale, the two Balkan species have small ranges. An early Pliocene time frame for the divergence of the subgenera Carinodreissena and Dreissena is discussed, as well as potential colonization routes of the most recent common ancestor of Carinodreissena spp. 5. The ambiguous taxonomy of dreissenids in the Balkans is addressed. As nominal D. blanci presbensis from Lake Prespa has nomenclatural priority over D. stankovici, the correct name for the latter taxon should be Dreissena presbensis.     

Molecular phylogenetics and biogeography of the Neotropical redstarts (Myioborus; Aves, Parulinae)

Montane areas in the Neotropics are characterized by high diversity and endemism of birds and other groups. The avian genus Myioborus (Parulinae) is a group of insectivorous warblers, characteristic of cloud forests, that represents one of the few Parulinae genera (New World warblers) that has radiated substantially in South America. The genus is distributed throughout most montane regions from the southwestern United States to northern Argentina. Here, I use mitochondrial sequences from the cytochrome b, ND2, and ND3 genes to present the first hypothesis of phylogenetic relationship among all Myioborus species level taxa. Phylogenetic reconstructions based on maximum parsimony, maximum likelihood, and Bayesian methods produced similar results and suggest a northern origin for the genus Myioborus with subsequent colonization of the Neotropical Montane Region. The lower-montane species, M. miniatus, is the sister taxon to a clade in which all taxa occupy upper-montane habitats. These "highland" taxa diverged early in the history of the genus and produced two well-defined monophyletic lineages, a Central-northern Andean clade formed by M. albifrons, M. ornatus, and M. melanocephalus, and a Pantepui (table-mountains of southern Venezuela, northern Brazil, and western Guyana) clade consisting of M. castaneocapillus, M. albifacies, and M. cardonai, and probably M. pariae. M. brunniceps, M. flavivertex, and M. torquatus were included in this upper-montane clade but without clear relationships to other taxa. Lack of resolution of nodes defining the upper-montane species clade is likely to result from a period of rapid diversification mediated by geological and climatic events during the Late Pliocene. These results suggest that an interplay of dispersal and vicariance has shaped the current biogeographic patterns of Myioborus.     

The influence of past and present climate on the biogeography of modern mammal diversity

Within most terrestrial groups of animals, including mammals, species richness varies along two axes of environmental variation, representing energy availability and plant productivity. This relationship has led to a search for mechanistic links between climate and diversity. Explanations have traditionally focused on single mechanisms, such as variation in environmental carrying capacity or evolutionary rates. Consensus, though, has proved difficult to achieve and there is growing appreciation that geographical patterns of species richness are a product of many interacting factors including biogeographic history and biological traits. Here, we review some current hypotheses on the causes of gradients in mammal richness and range sizes since the two quantities are intimately linked. We then present novel analyses using recent datasets to explore the structure of the environment-richness relationship for mammals. Specifically, we consider the impact of glaciation on present day mammalian diversity gradients. We conclude that not only are multiple processes important in structuring diversity gradients, but also that different processes predominate in different places.     

Early Hominid diversity, age and biogeography of the Malawi-Rift

Remains of earlyHomo andParanthropus have been recovered from two contemporaneous sites (Uraha and Malema) in the “Hominid Corridor” in Northern Malawi (Chiwondo Beds). Faunal dating suggests an age of 2.5–2.3 Ma for both hominids. The two specimens, a mandible attributed toHomo rudolfensis (UR 501 from Uraha), and a maxillary fragment ofParanthropus boisci. (RC 911 from Malema) known only from eastern Africa, represent the southernmost known distribution of these taxa. The biogeographic significance of these hominids from the Malawi-Rift lay in their association with the eastern African endemic animal group. Biogeographic variation in south-eastern Africa may be linked to habitat change occurring due to climate change, with maximum change occurring around 2.5 Ma.