PATTERNS AND PROCESSES OF DIVERSIFICATION: SPECIATION AND HISTORICAL CONGRUENCE IN SOME NEOTROPICAL BIRDS

This paper documents congruence in geographical patterns of speciation for four clades of birds having taxa endemic to the same areas within the Neotropics. Two genera, Pionopsitta parrots and Selenidera toucans, corroborate a well known biogeographic disjunction in which taxa endemic to southern Central America and the Chocó region of northwestern South America are the sister-group to a radiation within the Amazon basin. These two genera, along with two lineages within the toucan genus Pteroglossus, also document a pattern of historical interrelationships for four well known areas of endemism within Amazonia: Guyanan + (Belém-Pará + (Inambari + Napo)). These generalized historical patterns are interpreted to have arisen via fragmentation (vicariance) of a widespread ancestral biota. A review of the paleogeographic evidence suggests that these vicariance events could have originated as a result of several different mechanisms operating at various times during the Cenozoic. The inference that diversification of the Neotropical biota is primarily the result of the most recent of these possible vicariance events, namely isolation within Quaternary forest refugia, is unwarranted, given present data. These patterns of historical congruence are also interpreted as direct evidence against the hypothesis that diversification of the forest biota was a consequence of parapatric differentiation along recently established ecological gradients.     

Phylogenetic revision of the Charis cleonus complex (Lepidoptera: Riodinidae)

Abstract.  A revision of the Charis cleonus group of Neotropical riodinid butterflies is presented, including discussions on the taxonomy, biogeography and biology of its species, and illustrations of the adults and male and female genitalia (where known) of all taxa. We recognize twenty-two species, including nineteen new species: C. ariquemes sp.n., C. brasilia sp.n., C. breves sp.n., C. cacaulandia sp.n., C. cuiaba sp.n., C. humaita sp.n., C. ipiranga sp.n., C. iquitos sp.n., C. ma sp.n., C. manicore sp.n., C. manu sp.n., C. matic sp.n., C. maues sp.n., C. negro sp.n., C. palcazu sp.n., C. rocana sp.n., C. santarem sp.n., C. tapajos sp.n and C. tefe sp.n. A cladistic analysis using thirty-six characters of male and female genitalia and external facies generated a single most parsimonious cladogram highlighting the existence of two clades, the major and cleonus subgroups. As each species largely occupies a parapatric range, this cladogram provides the opportunity to reassess historical relationships among areas of endemism in the Amazon basin.     

A checklist of Lecythidaceae in the upper Madeira River,Rondônia,Brazil with comments on diversity and conservation

The Lecythidaceae or Brazil Nut family are ubiquitous in non-flooded lowland forests of the Amazon, where they are indicative of well-preserved or little-disturbed habitats. A recent checklist of the Brazilian flora reported 10 genera and 119 species for that country, of which 104 are found in the Amazon region. However, the botanical knowledge in many regions of the country remains far from complete. This study aimed to analyze the diversity of Lecythidaceae in the upper Madeira River region in Rondônia, an area where botanical data is scarce. Lecythidaceae collections deposited in the CEN, INPA and RON herbaria were examined, and four field expeditions in the study area were carried out. For that region, we recorded 37 species of Lecythidaceae belonging to 7 genera: Eschweilera (21 species), Couratari (6), Cariniana (3), Gustavia (3), Allantoma (2), Bertholletia (1) and Lecythis (1). Among these species, five are listed as threatened in the IUCN Red List. The total number of species found in the upper Madeira River of Rondônia alone is higher than the 28 species reported to date for the entire state of Rondônia in the Brazilian Flora Checklist (Smith et al., 2015), and adds 13 new records of Lecythidaceae for the state. Considering that this study is restricted to the northern part of the state, our results suggest that the number of Lecythidaceae species in Rondônia is severely underestimated. Our findings highlight the need for more intensive floristic studies in the Amazon region, which should target areas that are botanically unexplored but have high biodiversity potential. Such studies will provide valuable information to support both taxonomic studies and species conservation assessments.     

Fine-scale endemism of Amazonian birds in a threatened landscape

The region recognized as the Rondônia area of endemism is one of the most threatened in the Brazilian Amazon. In addition to rampant deforestation that took place during the 1980s and 1990s, it is now the locus of infrastructure development and economic expansion. However, the patterns of species distribution and diversity in this region are much more complex than previously recognized. Patterns of distribution of birds, primates, and butterflies strongly suggest that the Rondônia area of endemism is not biogeographically homogenous. On opposite banks of the Madeira tributaries, in particular the Aripuanã and Jiparaná rivers, there are genetically, morphologically, and vocally distinct bird populations with restricted distributions and geographic substitution. These areas are relatively small and already highly threatened by deforestation and economic development. Up to 70 % of the Jiparaná basin has been deprived of its original forest cover. Thus, their unique populations deserve to be recognized as conservation priorities and urgently included in conservation planning blueprints. Many restricted taxa, including many yet to be described, may already be highly endangered or locally extinct. Conservation planning in the Brazilian Amazon directed at specifying areas to be protected has focused on the larger proposed areas of endemism. If fine-scale endemism in threatened areas does exist in other Amazonian regions, then conservation policies that do not recognize these patterns will not adequately protect regional diversity.     

A study of Culicoides in Rondônia,in the Brazilian Amazon: species composition,relative abundance and potential vectors

There is very little information available about Culicoides species (Diptera: Ceratopogonidae) in the western Brazilian Amazon. However, studies of the fauna of this region are essential to knowledge of the species and potential vectors within it. Thus, the present study aims to evaluate the abundance, richness and composition of Culicoides species in rural areas in the state of Rondônia, Brazil. Culicoides specimens were collected in forest and pasture environments in the municipality of Porto Velho, using light traps. A total of 1708 individuals (1136 females and 572 males) belonging to 33 species were collected; 28 of these samples represent new records for the state of Rondônia and include the first record of Culicoides contubernalis in Brazil. Culicoides insignis was the most abundant species (86.1%). Species richness was greater in forest areas (32 species, 96.96%), whereas pastures presented the greatest number of Culicoides captured (n = 1540, 90.1%). This study shows that Culicoides populations differ between forest and pasture environments and indicates that the abundance of C. insignis is an important factor in epidemiological vigilance studies in the region.     

Integrating spatially explicit habitat projections into extinction risk assessments: a reassessment of Amazonian avifauna incorporating projected deforestation

Aim We aimed to complete the first systematic assessment of extinction risk based on projected population declines derived from spatially explicit habitat projections for any taxonomic group at a regional scale, to use the outputs to ascertain the efficacy of an existing protected area network in covering species of conservation concern, and identify gaps therein. Location This study focused on Amazonia; an area of exceptional biodiversity, currently experiencing the highest absolute rate of forest loss globally but where the proportion of species assessed as ‘threatened’ on the International Union for the Conservation of Nature (IUCN) Red List in the region is below global averages. Methods For all forest‐dependent Amazonian bird species (814), we revised extinction risk estimates by combining data from a spatially explicit deforestation model with generation length estimates. By overlaying distribution maps for these revised threatened species, we identified crisis areas (areas of projected deforestation supporting the highest numbers of threatened species), refugia (areas projected to retain forest supporting the highest numbers of threatened species) and areas of high irreplaceability: short‐ and long‐term priorities for new protected areas (PAs). Results The number of species qualifying as threatened rose substantially from 24 (3%) to 64–92 (8–11%). Areas of particular concern are the crisis and highly irreplaceable areas within the ‘arc of deforestation’ in the southern Brazilian Amazon states of Rondônia, Mato Grosso and Pará. Main conclusions Through a novel application of the IUCN Red List criteria, we present a spatially accurate rendering of the extinction risks of Amazonian birds. Important areas in the Amazon are not secure. We identify priorities for expansion of the PAs network and key locations where protection should be enforced. We recommend a collaborative approach employing our methods to repeat this process for other taxonomic groups.     

Genetic Diversity and Natural Population Structure of Cacao (Theobroma cacao L.) from the Brazilian Amazon Evaluated by Microsatellite Markers

A sample of 94 accessions of Theobroma cacao L. (cacao), representing four populations from the Brazilian Amazon (Acre, Rondônia, lower Amazon and upper Amazon) were analyzed using microsatellite markers to assess the genetic diversity and the natural population structure. From the 19 microsatellite loci tested, 11 amplified scorable products, revealing a total of 49 alleles, including two monomorphic loci. The Brazilian upper Amazon population contained the largest genetic diversity, with the most polymorphic loci, the highest observed heterozygosity; and the majority of rare alleles, thereby this region might be considered part of the center of diversity of the species. The observed heterozygosity for all the Brazilian populations (H _o = 0.347) was comparable with values reported for other similar upper Amazon Forastero cacao populations, with the Acre and Rondônia displaying the lowest values. The lower Amazon population, traditionally defined as highly homozygous, presented an unexpectedly high observed heterozygosity (H _o = 0.372), disclosing rare and distinct alleles, with large identity with the upper Amazon population. It was hypothesized that part of the lower Amazon population might derive from successive natural or intentional introduction of planting material from other provenances, mainly upper Amazon. Most of the loci exhibited a lower observed heterozygosity than expected, suggesting that self-pollination might be more common than usually assumed in cacao, but excess of homozygotes might also derive from sub-grouping (Wahlund effect) or from sampling related individuals. Most of the gene diversity was found to occur within groups, with small differentiation between the four Brazilian Amazon populations, typical of species with high gene flow.     

Campylocentrum benellii and C. paludosum spp. nov. (Angraecinae–Orchidaceae): two new leafless species from Brazil

The leafless condition is present in two neotropical epiphyte genera of Orchidaceae, Campylocentrum and Dendrophylax. Only the first one is reported from Brazil, where seven leafless species are recorded, and four of them are endemic. This study describes two new species of Campylocentrum from the Atlantic forest of São Paulo state and Amazon forest of Acre, Mato Grosso and Rondônia states. It also provides a key to the Brazilian species of leafless Campylocentrum, illustrations, and a discussion of the affinities of the new species with allied ones.     

Dung Beetle Communities and Seed Dispersal in Primary Forest and Disturbed Land in Amazonia1

Seeds from tropical fruiting trees ingested and defecated on the soil surface by primary dispersers (such as primates) are vulnerable to destruction from rodents, insects, and fungi. Burial by dung beedes as an incidental result of their feeding and nesting activities often provides these seeds with refugia from attack. To examine the effect of habitat disturbance on the dung beetle communities involved in this process, I surveyed dung beedes at three sites in the Amazon basin, in die states of Parí, Amazonas, and Rondónia, Brazil. Through principal component analysis on measurements of size and behavioral characters of beedes, I determined the relative quality of beetle species as seed dispersers (dispersal defined as horizontal or vertical movement of seeds) and ranked them into seed dispersal guilds. I used correspondence analysis to examine in what habitats (primary forest or varying degrees of disturbed habitat) these guilds were abundant. Most guilds decreased with increasing habitat disturbance, but one guild made up of large nocturnal burrowers (primarily Dichotomius) became more abundant with increasing disturbance (up to the level of highest disturbance surveyed), at which point all dung beede species became scarce. Clear–cuts had lower species richness, lower abundance, and lower biomass dian forested areas. These results imply that clear–cutting is detrimental to all seed dispersal dynamics in tropical rain forests, but that some levels of disturbance allow enough disperser activity to preserve this ecosystem function.     

New and interesting species of Hymenochaetaceae from the Brazilian Amazonia

Coltricia globispora sp. nov. is described as new based on specimens collected in the Parque Natural Municipal de Porto Velho, in the state of Rondônia, Brazilian Amazonia. It is characterized by small pores (7–8 per mm), monomitic hyphal system, and globose, smooth and guttulate basidiospores (5.5–6.6 (?7) μm). Phellinus griseoporus has been found for the first time in Brazil, and Cyclomyces iodinus, P. merrillii, P. membranaceus and P. umbrinellus in the Brazilian Amazonia. Coltricia cinnamomea, P. calcitratus and Phylloporia chrysites are new occurrences to the state Rondônia, while C. hamata and Phylloporia spathulata to the state of Pará. The knowledge about Hymenochaetaceae in the Brazilian Amazonia is briefly discussed, and a key to the neotropical species of Coltricia is provided.     

Historical biogeography of New World emballonurid bats (tribe Diclidurini): taxon pulse diversification

Aim To reconstruct the biogeographical history of New World emballonurid bats (tribe Diclidurini). Although bats are the second most species‐rich order of mammals, they have not contributed substantially to our understanding of the historical biogeography of mammals in the Neotropics because of a poor fossil record. In addition, being the only group of mammals that fly, bats typically have large distributions with relatively few species endemic to restricted areas that are amenable to vicariant biogeographical approaches. Location Central and South America. Methods Phylogenetic analysis for comparing trees (PACT) is a new algorithm that incorporates all spatial information from taxon area cladograms into a general area cladogram. There were nine biogeographical areas identified in Central and South America for New World emballonurid bats. Molecular dating was used to incorporate the temporal aspect of historical biogeography. This method was compared with dispersal–vicariance analysis (DIVA), which assumes vicariance as the default mode of speciation. Results Of the 45 speciation events in a fully resolved phylogeny, eight that were hypothesized by DIVA as vicariance were considered by PACT as two peripheral isolations and six within‐area events. DIVA was less parsimonious because it required six more post‐speciation dispersal events in addition to the 73 hypothesized by PACT. DIVA reconstructed a widely distributed ancestor, suggesting that most dispersal events occurred earlier, whereas the ancestral area for PACT based on character optimization was the Northern Amazon, suggesting that dispersal events were more recent phenomena. Main conclusions The general area cladogram from PACT indicated that within‐area events, and not vicariance, provide the major mode of speciation for New World emballonurid bats. There was no biological evidence supporting or rejecting sympatric speciation in New World emballonurid bats. However, the geological history, combined with fluctuations in temperature and sea level, suggested within‐area speciation in a changing and heterogeneous environment in the Northern Amazon during the Miocene. This scenario is similar to the taxon‐pulse hypothesis of biotic diversification, which posits repeated episodes of range expansions and contractions from a stable core area such as the Guiana Shield within the Northern Amazon.     

Tree Species Distribution along Environmental Gradients in an Area Affected by a Hydroelectric Dam in Southern Amazonia

We describe the patterns of composition and abundance of tree species and their relationships to environmental factors in an area affected by the Jirau hydroelectric dam on the Madeira River in the Brazilian state of Rondônia. Trees were sampled in 20 1‐ha forest plots distributed among areas affected and unaffected by the hydroelectric reservoir. Predictors of species distribution included in the analyses were soil fertility (sum of bases) and texture, slope, and the vertical distance from the nearest drainage. We sampled 8504 individuals belonging to 909 species/morphospecies. Floristic differences occurred throughout the reservoir area and between the sides of the river, and were strongly associated with edaphic and geological factors. The predictor with the greatest effect on the floristic changes was the sum of bases, coupled with the vertical distance from the nearest drainage, the latter reflecting the influence of the riparian zone. The forests sampled were heterogeneous and included transitional formations between unflooded terra firme and seasonally inundated (varzea, campinarana) forests. Although a considerable number of species (253) were restricted to plots affected by the reservoir, most of these species have a widespread distribution in the Amazon region. However, conservation measures are required in order to preserve species associations and ecosystems processes similar to those most affected by the reservoir such as the river floodplains. Our study reinforces the need to consider the heterogeneity of habitats affected by reservoirs when making decisions concerning the building of hydroelectric dams in the Amazon.     

Soil organic carbon stock change due to land use activity along the agricultural frontier of the southwestern Amazon,Brazil, between 1970 and 2002

The southwestern portion of the Brazilian Amazon arguably represents the largest agricultural frontier in the world, and within this region the states of Rondônia and Mato Grosso have about 24% and 32% of their respective areas under agricultural management, which is almost half of the total area deforested in the Brazilian Amazon biome. Consequently, it is assumed that deforestation in this region has caused substantial loss of soil organic carbon (SOC). In this study, the changes in SOC stocks due to the land use change and management in the southwestern Amazon were estimated for two time periods from 1970–1985 and 1985–2002. An uncertainty analysis was also conducted using a Monte Carlo approach. The results showed that mineral soils converted to agricultural management lost a total of 5.37 and 3.74 Tg C yr^?1 between 1970–1985 and 1985–2002, respectively, along the Brazilian Agricultural Frontier in the states of Mato Grosso and Rondônia. Uncertainties in these estimates were ±37.3% and ±38.6% during the first and second time periods, respectively. The largest sources of uncertainty were associated with reference carbon (C) stocks, expert knowledge surveys about grassland condition, and the management factors for nominal and degraded grasslands. These results showed that land use change and management created a net loss of C from soils, however, the change in SOC stocks decreased substantially from the first to the second time period due to the increase in land under no‐tillage.     

Epidemiological and nosological aspects of Leishmania naiffi Lainson & Shaw, 1989.

Leishmania naiffi was isolated from 10 out of 64 armadillos (Dasypus novemcinctus) examined in Amazonas, Pará and Rond?nia States in the Brazilian Amazon Region. The isolates were obtained in culture from samples of liver (3), spleen (3), lymph nodes (2), skin (1) and blood (1) from the infected animals. Heavy infections with the same parasite were detected for the first time in Psychodopygus squamiventris, a common man-biting phlebotomine, in Amazonas and Pará. A new case of cutaneous leishmaniasis caused by L. naiffi is described from the Manaus area, making a total of three known cases of human infection by this parasite.     

Geographic distribution and abundance of woolly (Lagothrix cana) and spider (Ateles chamek) monkeys in southwestern Brazilian Amazonia.

Two species of frugivorous atelids, Ateles chamek and Lagothrix cana, occur in southwestern Brazilian Amazonia. Populations were surveyed at 36 sites in the state of Rondônia. Ateles chamek is widespread, but the distribution of L. cana is limited by a combination of riverine barriers and ecological factors, possibly including competition with A. chamek. Groups of L. cana were generally larger and more abundant than those of A. chamek, even in syntopy. The transitional forest that predominates in the extreme south of Rondônia (Hylea‐cerrado) is not a barrier to either species, with both species being tolerant of habitat disturbance when hunting pressure is low. Am. J. Primatol. 56:57–64, 2002. © 2002 Wiley‐Liss, Inc.     

Using indirect methods to constrain symbiotic nitrogen fixation rates: a case study from an Amazonian rain forest

Human activities have profoundly altered the global nitrogen (N) cycle. Increases in anthropogenic N have had multiple effects on the atmosphere, on terrestrial, freshwater and marine ecosystems, and even on human health. Unfortunately, methodological limitations challenge our ability to directly measure natural N inputs via biological N fixation (BNF)—the largest natural source of new N to ecosystems. This confounds efforts to quantify the extent of anthropogenic perturbation to the N cycle. To address this gap, we used a pair of indirect methods—analytical modeling and N balance—to generate independent estimates of BNF in a presumed hotspot of N fixation, a tropical rain forest site in central Rondônia in the Brazilian Amazon Basin. Our objectives were to attempt to constrain symbiotic N fixation rates in this site using indirect methods, and to assess strengths and weaknesses of this approach by looking for areas of convergence and disagreement between the estimates. This approach yielded two remarkably similar estimates of N fixation. However, when compared to a previously published bottom-up estimate, our analysis indicated much lower N inputs via symbiotic BNF in the Rondônia site than has been suggested for the tropics as a whole. This discrepancy may reflect errors associated with extrapolating bottom-up fluxes from plot-scale measures, those resulting from the indirect analyses, and/or the relatively low abundance of legumes at the Rondônia site. While indirect methods have some limitations, we suggest that until the technological challenges of directly measuring N fixation are overcome, integrated approaches that employ a combination of model-generated and empirically-derived data offer a promising way of constraining N inputs via BNF in natural ecosystems.     

Historical biogeography of some river basins in central Mexico evidenced by their goodeine freshwater fishes: a preliminary hypothesis using secondary Brooks parsimony analysis

Aims Our aim was to uncover and describe patterns of historical biogeography of the main river basins in central Mexico, based on a secondary Brooks parsimony analysis (BPA) of goodeine fishes, and to understand the processes that determine them with respect to the molecular clock of the goodeines and the geological events that have taken place in the region since the Miocene. Location The region covered in this study includes central Mexico, mostly the so‐called Mesa Central of Mexico, an area argued to be a transitional zone comprising several major river drainages from their headwaters at high elevations along the Transmexican Volcanic Belt to the coast of the Gulf of Mexico and the Pacific Ocean. Methods Based on a previous phylogenetic hypothesis regarding the Goodeidae, we built a data matrix using additive binary coding. First, we conducted a primary BPA to provide general explanations of the historical biogeography of Central Mexico. As ambiguity was found, a secondary BPA was conducted, and some areas were duplicated in order to explain the reticulated history of the area. Area cladograms were obtained by running a parsimony analysis. Instances of vicariance and non‐vicariance processes were described with reference to the cladogram obtained from secondary BPA. Results The study area was divided into 18 discrete regions. Primary BPA produced nine equally parsimonious cladograms with 129 steps, and a consistency index (CI) of 0.574. A strict consensus cladogram shows low resolution among some areas, but other area relationships are consistent. For secondary BPA, five of the 18 regions were duplicated (LEA, COT, AYU, CUT, PAN); one was triplicated (BAL); and one was quadruplicated (AME), suggesting that the pattern of distribution of species in these areas reflects multiple independent events. These areas correspond with the regions exhibiting the highest levels of diversification and the most complex geological history, and those for which river piracy events or basin connections have been proposed. The secondary BPA produced a single most parsimonious cladogram with 118 steps, and a CI of 0.858. This cladogram shows that none of the duplicated areas are nested together, reinforcing the idea of a reticulated history of the areas and not a single vicariant event. Main conclusions Although our results are preliminary and we cannot establish this as a general pattern, as the BPA is based on a single‐taxon cladogram, resolution obtained in the secondary BPA provides some insights regarding the historical biogeography of this group of fishes in river basins of central Mexico. Secondary BPA indicates that the historical biogeography of central Mexico, as shown by their goodeine freshwater fishes, is complex and is a result of a series of vicariant and non‐vicariant events such as post‐dispersal speciation and post‐speciation dispersal.     

New foci of transmission of Schistosoma mansoni in the state of Pará

Two new foci of transmission of Schistosoma mansoni in the state of Pará are recorded, with the finding of naturally infected Biomphalaria glabrata in the municipalities of Viseu and Belém. Uninfected specimens of Biomphalaria straminea, as well as the planorbid species Biomphalaria schrammi, Drepanotrema lucidum and D. anatinum, were found in the same area.     

Comments on hexanchiform phylogeny (Pisces, Neoselachii)

The Hexanchiformes (Cow Sharks) are regarded as a monophyletic taxon. Cladistic analysis shows that among the various neoselachian taxa proposed so far as the sister group of the Hexanchiformes a sister group relationship between the Hexanchiformes and a (still unnamed) taxon comprising the Squaliformes and Pristiophoriformes appears as the most probable hypothesis. In addition, MAISEY and WOLFRAM'S (1984) concept of hexanchiform interrelationships is critically reviewed. An alternative cladogram of hexanchiform interrelationships is developed which includes Recent as well as fossil hexanchiform taxa. In this cladogram the living genera Hexancbus and Notorynchus are sister groups and both taxa together form the sister group of the Recent Heptranchias. The fossil taxa +Notidanoides, +“Hexanchus” gracilis, +Notidanodon and +Weltonia are arranged in the stem lineage of recent Hexanchiformes.     

PACT: an efficient and powerful algorithm for generating area cladograms

Aim To introduce and describe the functioning of a new algorithm, phylogenetic analysis for comparing trees (PACT), for generating area cladograms that provide accurate representation of information contained in taxon–area cladograms. Methods PACT operates in the following steps. Convert all phylogenies to taxon–area cladograms. Convert all taxon–area cladograms to Venn diagrams. Choose any taxon–area cladogram from the set of taxon–area cladograms to be analysed and determine its elements. This will be the template area cladogram. Select a second taxon–area cladogram. Determine its elements. Document which elements in the second tree occur in the template tree (denoted by ‘Y’) and which do not (denoted by ‘N’). Each ‘Y’ indicates a match with previous pattern and these are combined. Each ‘N’ is a new element and is attached to the template area cladogram at the node where it is linked with a Y. This requires two rules: (1) ‘Y + Y = Y’ (combine common elements) as long as they are connected at the same node; and (2) ‘Y + N = YN’ (add novel elements to the template area cladogram at the node where they first appear). Once the novel elements in the second taxon–area cladogram have been added to the template area cladogram, see if any of them can be further combined. This requires three additional rules: (1) ‘Y(Y? = Y(Y?’ (do not combine Y's if they are attached at different nodes on the template area cladogram); (2) ‘Y + YN = YN’ (Y is part of group YN); and (3) ‘YN + YN = YNN’ (Y is the same for each, but each N is different). Repeat for all available taxon–area cladograms. Results Three exemplars demonstrate that PACT provides the most accurate area cladograms for vicariance‐driven biotic diversification, dispersal‐driven biotic diversification and taxon pulse‐driven biotic diversification. PACT can also be used as an a priori method of biogeographical analysis. Main conclusions PACT embodies all the strong points and none of the weaknesses of previously proposed methods of historical biogeography. It is most useful as an a posteriori method, but it is also superior to all previous a priori methods because it does not specify costs, or weights or probabilities, or likelihoods of particular biogeographical processes a priori and is thus sensitive to clade‐specific historical contingencies.