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Identifying global zoogeographical regions: Lessons from Wallace

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Abstract

Aim When dividing the world into zoogeographical regions, Alfred Russel Wallace stipulated a set of criteria by which regions should be determined, foremost the use of generic rather than species distributions. Yet, recent updates of Wallace’s scheme have not followed his reasoning, probably explain- ing in part the discrepancies found. Using a recently developed quantitative method, we evaluated the world’s zoogeographical regions following his criteria as closely as possible. Location Global. Methods We subjected presence–absence data from range maps of birds, mammals and amphibians to an innovative clustering algorithm, affinity prop- agation. We used genera as our taxonomic rank, although species and familial ranks were also assessed, to evaluate how divergence from Wallace’s criteria influences the results. We also accepted Wallace’s argument that bats and migratory birds should be excluded (although he was contradictory about the birds) and devised a procedure to determine the optimal number of regions to eliminate subjectivity in delimiting the number of regions. Results Regions attained using genera (eight for mammals and birds and six for amphibians) strongly coincided with the regions proposed by Wallace. The regions for amphibians were nearly identical to Wallace’s scheme, whereas we obtained two new ‘regions’ for mammals and two for birds that largely coin- cide with Wallace’s subregions. As argued by Wallace, there are strong reasons not to consider these as being equivalent to the six main regions. Species distri- butions generated many small regions related to contemporary climate and vegetation patterns, whereas at the familial rank regions were very broad. The differences between our generic maps and Wallace’s all involve areas which he identified as being uncertain in his regionalization. Main conclusions Despite more than 135 years of additional knowledge of distributions, the shuffling of generic concepts, and the development of com- puters and complex analytical techniques, Wallace’s zoogeographical regions appear to be no less valid than they were when he proposed them. Recent studies re-evaluating Wallace’s scheme should not be considered updates as such because they have not followed Wallace’s reasoning, and all computer- based analyses, including this one, are subject to the vagaries of the particular methods used.

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... In the last decade or so, several research teams have re-visited the global biogeographical realms issue, and in doing so have delineated patterns in the Indo-Australian Archipelago (e.g. Holt et al., 2013;Kreft & Jetz, 2010;Procheş & Ramdhani, 2012;Rueda et al., 2013;Figure 1c). Relative to the 19th Century workers, the groups were able to draw upon vastly improved taxa-occurrence records and to apply modern-day analytical techniques. ...
... Relative to the 19th Century workers, the groups were able to draw upon vastly improved taxa-occurrence records and to apply modern-day analytical techniques. Kreft and Jetz (2010), Holt et al. (2013) and Rueda et al. (2013) It is largely the same as Merrill's (1924) Wallacea, but they treated it as equivalent in standing to the adjacent Palearctic, Indo-Malaysian, New Guinean and F I G U R E 4 Maps of the Indo-Australian archipelago and adjacent areas showing various biogeographical divides and areas that are mentioned in the text. Panel (a) depicts Wallacea as originally defined by Merrill (1924), the 'reduced' version that is widely used today (which was introduced by Darlington, 1957), plus the Wallacea linguistics area (e.g. ...
... Schapper, 2015). Panel (b) portrays the key lines and area that were proposed in the last decade or so by Kreft and Jetz (2010), Procheş & Ramdhani, 2012, Holt et al. (2013 and Rueda et al. (2013). The base image was generated using GeoMapApp (Ryan et al., 2009). ...
Article
To mark A.R. Wallace's 200th birthday, we review the direct and indirect contributions he made to our understanding of the Indo‐Australian Archipelago's biogeography. He is widely known for his field research (1854–1862) and his 1863 boundary line separating the Oriental and Australasian faunal realms (between Bali and Lombok, Borneo and Sulawesi, and the Philippines and the Moluccas). Notably, though, he never accepted Huxley's ‘Wallace Line’ proposal (1868), whose northern part runs between the main Philippine islands and the Palawan Group to the west. Furthermore, in 1910, which was 3 years prior to his demise, he transferred Sulawesi's fauna to the Oriental realm. In 1924, Merrill introduced the ‘Wallacea’ transition zone. Although the label is today widely used to denote a sub‐region within the Indo‐Australian Archipelago between Wallace's 1863 line and Lydekker's 1896 line (first presented by Darlington in 1957), the western boundary was originally based on Huxley's line, and thus included the Philippine islands minus the Palawan group. Most biogeographers appear to be unaware of Merrill and his intention. Finally, recent attempts to define the faunal break have not led to a consensus view, despite the huge increase in primary data plus the application of modern analytical techniques. This reflects the complexities and diversity of the region's faunal distribution patterns, plus the differences in the ways that researchers choose to process their data.
... (o) Morrone (2015) Responding to the various global regionalization studies that had been published in the previous 5 years (e.g. Kreft & Jetz, 2010;Procheş & Ramdhani, 2012;Holt et al., 2013;Rueda et al., 2013), Morrone (2015) proposed a qualitative compromise. For the Indo-Australian Archipelago he inserted into the eastern end of his Oriental Region (Morrone, 2002; see Section III.3b) a transition zone that replicated Darlington (1957), that is, 'reduced' Wallacea. ...
... In the last decade new lines have been proposed as a consequence of quantitative analyses (e.g. Kreft & Jetz, 2010;Holt et al., 2013;Rueda et al., 2013; see also Lohman et al., 2011), or the integration of new data (Ali et al., 2020). ...
... Concerning modern studies that have applied state-of-theart, quantitative analyses to this region (e.g. Kreft & Jetz, 2010;Procheş & Ramdhani, 2012;Holt et al., 2013;Rueda et al., 2013), the inescapable fact is that we are no nearer to reaching agreement on where any lines/areas should be placed than we were more than a century ago. Given the fact that Wallacea contains a large and diverse set of organisms, each with its own history, this is not unexpected. ...
Article
Due to its position between the highly distinct Oriental and Australasian biogeographical realms, much effort has been spent demarcating associated separations and transitions in the faunal assemblages of the Indo‐Australian Archipelago. Initially, sharp boundary lines were proposed, with the earliest dating from the mid‐1800s. Notably, the one published by Alfred R. Wallace in 1863, based upon land‐mammal and land‐bird distributions, has since achieved iconic status and today its significance is recognized well beyond the confines of the biogeography community. Over the next four decades many such divides were engraved onto plates and inked onto charts of SE Asia using additional information, different organisms or other criteria. However, it became apparent that, as Wallace had noted, all such lines were to some degree permeable, and by the 1880s transition zones were being put forward instead; the label ‘Wallacea’ was introduced in 1924. Interestingly, the last decade has seen new divides and sub‐regions being proposed, some departing markedly from earlier offerings. Although currently there is general agreement regarding much of the terminology associated with both the lines and the areas, the record of publication indicates that this consensus has emerged obliquely, and in some cases is weakly founded. This review does not present new data nor new analyses; rather it summarizes the development of ideas and reflects upon attendant issues that have emerged. After reviewing the key proposals, recommendations are presented that should in future alleviate perceived difficulties or inadequacies. Reference to specific divides must be true to their original definitions; there are many instances where the secondary literature has portrayed them incorrectly and with some this has rippled through into later publications. Moreover, Wallace's 1863 line is not the one that he finally settled upon (in 1910); its path around Sulawesi was transferred from the west to the east of the Island. Ideally, Huxley's divide (1868) should carry his name rather than Wallace's; the latter never accepted the proposition. Lydekker's Line (1896) ought to be labelled the Heilprin–Lydekker Line in recognition of Angelo Heilprin's 1887 contribution. Concerning transition zones, ideally Wallacea should correspond to its original 1924 description, which incorporated the Philippine islands bar the Palawan group. Notably, though, a smaller form (introduced by Darlington in 1957, used frequently from 1998 onwards) in which all of the Philippine islands are excluded is entrenched within the recent literature, but this is often without evident justification. It should also be recognized that the ‘reduced’ (=southern) Wallacea area was effectively defined by Heilprin in 1887, but was then labelled the ‘Austro‐Malaysian Transition Zone’. Finally, the application in recent years of modern analytical techniques has not led to a consensus view on where the lines/areas should run/be placed; with such a large, diverse set of organisms, each with differing histories, this is perhaps not surprising.
... We estimated the total number of species in each clade and genus occurring in each of these regions. We defined the Nearctic, Neotropical, and Austral biogeographical regions according to the Sclater-Wallace system [16,75] considering the modifications for bird genera proposed by Rueda et al. [75]. The Nearctic region extends from Alaska to the Trans-Mexican Volcanic Belt (around 19˚to 20˚N). ...
... We estimated the total number of species in each clade and genus occurring in each of these regions. We defined the Nearctic, Neotropical, and Austral biogeographical regions according to the Sclater-Wallace system [16,75] considering the modifications for bird genera proposed by Rueda et al. [75]. The Nearctic region extends from Alaska to the Trans-Mexican Volcanic Belt (around 19˚to 20˚N). ...
... The Neotropical region includes the tropical lowlands adjacent to the Mexican highlands and extends southwards, including the northern and central portions of South America. Finally, the Austral region comprises the southern Andes from Peru to the Patagonia (Fig 1C in Rueda et al. [75]). ...
Article
Full-text available
Mutualistic interactions are powerful drivers of biodiversity on Earth that can be represented as complex interaction networks that vary in connection pattern and intensity. One of the most fascinating mutualisms is the interaction between hummingbirds and the plants they visit. We conducted an exhaustive search for articles, theses, reports, and personal communications with researchers (unpublished data) documenting hummingbird visits to flowers of nectar-rewarding plants. Based on information gathered from 4532 interactions between 292 hummingbird species and 1287 plant species, we built an interaction network between nine hummingbird clades and 100 plant families used by hummingbirds as nectar resources at a continental scale. We explored the network architecture, including phylogenetic, morphological, biogeographical, and distributional information. As expected, the network between hummingbirds and their nectar plants was heterogeneous and nested, but not modular. When we incorporated ecological and historical information in the network nodes, we found a generalization gradient in hummingbird morphology and interaction patterns. The hummingbird clades that most recently diversified in North America acted as generalist nodes and visited flowers with ornithophilous, intermediate and non-ornithophilous morphologies, connecting a high diversity of plant families. This pattern was favored by intermediate morphologies (bill, wing, and body size) and by the low niche conservatism in these clades compared to the oldest clades that diversified in South America. Our work is the first effort exploring the hummingbird-plant mutualistic network at a continental scale using hummingbird clades and plant families as nodes, offering an alternative approach to exploring the ecological and evolutionary factors that explain plant-animal interactions at a large scale.
... Global regionalisation after 1946 follows a similar theme of that of later 19th century area classification, including how to reconcile the Holarctic with the Nearctic and Palaearctic regions. Late 20th and 21st century biogeographical regionalisation may be divided into the following three groups: those who, knowingly or unknowingly, rejected the Holarctic in favour of the Nearctic and Palearctic Kreft and Jetz 2010;Procheş and Ramdhani 2012;Holt et al. 2013aHolt et al. , 2013bRueda et al. 2013); those who accepted only a unified Holarctic (e.g. de Lattin 1967;; and those who attempted to incorporate each under a larger www.publish.csiro.au/sb Australian Systematic Botany ...
... Wildfinder data, both of these approaches tended to yield clusters of very uneven geographic coverage, often small clusters in tropical America and large Holarctic or Palaeotropical clusters [Procheş and Ramdhani 2012, p. 261]. Rueda et al. (2013) recovered regions similar to those of Wallace (1876a, 1876b), using amphibian, bird and mammal data, and Escalante (2017) recovered Wallace's Australian, Ethiopian, Neotropical, and Oriental regions by using mammal data. Neither study found evidence for the Holarctic. ...
... Nearctic region: Sclater (1858, p. 142); Kirby (1872, p. 437); Wallace (1876b, p. 114); Sharpe (1884, p. 102); Heilprin (1887, p. 62); Sclater (1894, p. 98); Sclater and Sclater (1899, p. 153); Bartholomew et al. (1911, p. 10); Darlington (1957, p. 442); Rapoport (1968, p. 94); Morrone ( , p. 150, 2006); Procheş and Ramdhani (2012, p. 263); Kreft and Jetz (2013, p. 343-c); Rueda et al. (2013Rueda et al. ( , p. 2217; Ribeiro et al. (2014, p. 249); Morrone (2015a, p. 84); Escalante (2017, p. 357); Hermogenes de Mendonça and Ebach (2020, p. 721); Escalante et al. (2021, p. 354). ...
Article
Full-text available
An interim hierarchical classification (i.e. biogeographical regionalisation or area taxonomy) of the world’s terrestrial regions is provided, following the work of Morrone published in Australian Systematic Botany in 2015. Area names are listed according to the International Code of Area Nomenclature so as to synonymise redundant names. The interim global terrestrial regionalisation to the subregion level recognises 3 kingdoms, 2 subkingdoms, 8 regions, 21 subregions and 5 transition zones. No new names are proposed for the regions; however, Lydekker’s Line is renamed Illiger’s Line. We note that some regions still require area classification at the subregion level, particularly the Palearctic, Ethiopian and Oriental regions. Henceforth, the following interim global regionalisation may be used as a template for further revisions and additions of new areas in the future.
... This upsurge was facilitated by the increase in quality and quantity of large-scale datasets, as well as the development of new analytical tools (Edler, Guedes, Zizka, Rosvall, & Antonelli, 2016;Kreft & Jetz, 2010;Vilhena & Antonelli, 2015). Consequently, multiple studies have tried to identify the major biogeographical regions for birds (Holt et al., 2013;Procheş & Ramdhani, 2012;Rueda, Rodríguez, & Hawkins, 2013), mammals (Holt et al., 2013;Kreft & Jetz, 2010;Procheş & Ramdhani, 2012;Rueda et al., 2013), amphibians (Edler et al., 2016;Holt et al., 2013;Procheş & Ramdhani, 2012;Rueda et al., 2013;Vilhena & Antonelli, 2015) and reptiles (Procheş & Ramdhani, 2012). The result of this upsurge was a debate on the precise limits of biogeographical regions. ...
... This upsurge was facilitated by the increase in quality and quantity of large-scale datasets, as well as the development of new analytical tools (Edler, Guedes, Zizka, Rosvall, & Antonelli, 2016;Kreft & Jetz, 2010;Vilhena & Antonelli, 2015). Consequently, multiple studies have tried to identify the major biogeographical regions for birds (Holt et al., 2013;Procheş & Ramdhani, 2012;Rueda, Rodríguez, & Hawkins, 2013), mammals (Holt et al., 2013;Kreft & Jetz, 2010;Procheş & Ramdhani, 2012;Rueda et al., 2013), amphibians (Edler et al., 2016;Holt et al., 2013;Procheş & Ramdhani, 2012;Rueda et al., 2013;Vilhena & Antonelli, 2015) and reptiles (Procheş & Ramdhani, 2012). The result of this upsurge was a debate on the precise limits of biogeographical regions. ...
... This upsurge was facilitated by the increase in quality and quantity of large-scale datasets, as well as the development of new analytical tools (Edler, Guedes, Zizka, Rosvall, & Antonelli, 2016;Kreft & Jetz, 2010;Vilhena & Antonelli, 2015). Consequently, multiple studies have tried to identify the major biogeographical regions for birds (Holt et al., 2013;Procheş & Ramdhani, 2012;Rueda, Rodríguez, & Hawkins, 2013), mammals (Holt et al., 2013;Kreft & Jetz, 2010;Procheş & Ramdhani, 2012;Rueda et al., 2013), amphibians (Edler et al., 2016;Holt et al., 2013;Procheş & Ramdhani, 2012;Rueda et al., 2013;Vilhena & Antonelli, 2015) and reptiles (Procheş & Ramdhani, 2012). The result of this upsurge was a debate on the precise limits of biogeographical regions. ...
Article
To define the major biogeographical regions and transition zones for freshwater fish species. Strictly freshwater species of actinopterygian fish (i.e. excluding marine and amphidromous fish families). We based our bioregionalization on a global database of freshwater fish species occurrences in drainage basins, which, after filtering, includes 11,295 species in 2,581 basins. On the basis of this dataset, we generated a bipartite (basin‐species) network upon which we applied a hierarchical clustering algorithm (the Map Equation) to detect regions. We tested the robustness of regions with a sensitivity analysis. We identified transition zones between major regions with the participation coefficient, indicating the degree to which a basin has species from multiple regions. Our bioregionalization scheme showed two major supercontinental regions (Old World and New World, 50% species of the world and 99.96% endemics each). Nested within these two supercontinental regions lie six major regions (Nearctic, Neotropical, Palearctic, Ethiopian, Sino‐Oriental and Australian) with extremely high degrees of endemism (above 96% except for the Palearctic). Transition zones between regions were of limited extent compared to other groups of organisms. We identified numerous subregions with high diversity and endemism in tropical areas (e.g. Neotropical), and a few large subregions with low diversity and endemism at high latitudes (e.g. Palearctic). Our results suggest that regions of freshwater fish species were shaped by events of vicariance and geodispersal which were similar to other groups, but with freshwater‐specific processes of isolation that led to extremely high degrees of endemism (far exceeding endemism rates of other continental vertebrates), specific boundary locations and limited extents of transition zones. The identified bioregions and transition zones of freshwater fish species reflect the strong isolation of freshwater fish faunas for the past 10–20 million years. The extremely high endemism and diversity of freshwater fish fauna raises many questions about the biogeographical consequences of current introductions and extinctions.
... The earliest of these studies focused on East Asian freshwater fish (Berg 1912(Berg , 1916Mori 1936), after which Chinese researchers studied the zoogeographic divisions of individual freshwater faunas at a national level (Zhang 1954;Li 1981;. More recently, the biogeographic divisions for freshwater fishes, amphibians and freshwater crabs, in China or globally, have all been variously studied individually (Chen & Bi 2007;Rueda et al. 2013;Holt et al. 2013;Kang et al. 2014;Vilhena & Antonelli 2015;Edler et al. 2016;Leroy et al. 2019). Freshwater ecoregions of the world have been proposed (Abell et al. 2008), but they are not necessarily based on species distributions and not quantitatively derived. ...
... These regions included: Southwest, Southeast, Northeast, Middle-East, Middle-Northwest and Hainan Island and Taiwan Island; however, the study largely lacked data from northern and western China (Fig. 10). The global zoogeographic studies by Holt (2013), Rueda et al. (2013), Vilhena & Antonelli (2015) and Edler et al. (2016) all included individual analyses on amphibians. Holt et al. (2013) found China to be part of three different amphibian zoogeographic regions while the areas found by Rueda et al. (2013), using genera occurrence data, resembled those of Wallace's (1876) regions. ...
... The global zoogeographic studies by Holt (2013), Rueda et al. (2013), Vilhena & Antonelli (2015) and Edler et al. (2016) all included individual analyses on amphibians. Holt et al. (2013) found China to be part of three different amphibian zoogeographic regions while the areas found by Rueda et al. (2013), using genera occurrence data, resembled those of Wallace's (1876) regions. Vilhena & Antonelli (2015) and Edler et al. (2016) both used a network approach and obtained areas with similar margins. ...
Article
Biogeographic regionalisations extract patterns of co-occurrence from different taxa to form a hierarchical system of geographical units of different scales. This system is useful for revealing biogeographic patterns and can be used as the basis for scientific communication between different fields. The history of Chinese freshwater biogeography is not well known to most modern biogeographers and is reviewed herein. We produce the first quantitative bioregionalisation of the freshwater zoogeographic areas of mainland China based on multiple animal groups. The combined occurrence data of amphibians, freshwater fish and freshwater crabs were subjected to cluster and network analyses. The two different methods yielded largely similar results. We propose four freshwater zoogeographical subregions (Beifang, Tarim, China, and the Tibetan subregion), three dominions for the China subregion (Jianghuai, Dongyang, and the new Dian dominion), three provinces for the Dian dominion (West Hengduan, Diannan Highlands and the new Yungui Plateau province) and two provinces for the Dongyang dominion (Zhemin and the new Huanan province) according to the naming rules of ICAN. The endemic areas of each animal group were then individually studied and were found to reflect the bioregionalisation at the subregion level, but differed from each other at the dominion and province level. Our analyses show that: (1) previous intuitive biogeographical studies have found similar areas; (2) there are recurring large scale biogeographic patterns in Chinese freshwater fishes, amphibians and freshwater crabs; and (3) bioregionalisations derived from quantitative methods can be effective for partitioning areas into biogeographically meaningful units.
... The biogeographic regionalisation of the world proposed by Wallace (1876) has been re-analysed several times (Udvardy 1975;Cox 2001;Procheş 2005;Kreft and Jetz 2010;Procheş and Ramdhani 2012;Holt et al. 2013;Rueda et al. 2013;Escalante et al. 2014;Morrone 2014aMorrone , 2015. These reanalyses have resulted in biogeographic regions that exhibit some similarities, but also differ from Wallace's classification. ...
... Udvardy (1975) considered a different northern boundary for the NR, namely Baja California and Sinaloa Coast, Mexico. Some studies agree that the NR extends from northern Mexico to central Argentina (Cabrera and Willink 1973;Smith 1983;Morrone 2001b;Kreft and Jetz 2010;Holt et al. 2013;Rueda et al. 2013). It has been proposed that the NR excludes the Andean region (Morrone 2004(Morrone , 2014a(Morrone , 2015Kreft and Jetz 2010;Procheş and Ramdhani 2012;Noguera-Urbano 2013;Noguera-Urbano and Escalante 2015), and that it includes the Mexican transition zone (North) and the South American transition zone (South; Morrone 2004Morrone , 2014aMorrone , 2015Noguera-Urbano 2013;Noguera-Urbano and Escalante 2015). ...
... Many protocols and algorithms have been described for the analysis of biogeographic areas (regions, subregions, provinces, ecoregions;Rosen 1988aRosen , 1988bRosen and Smith 1988;Harold and Mooi 1994;Morrone 1994;Hausdorf and Hennig 2003;Kreft and Jetz 2010;Holt et al. 2013;Rueda et al. 2013), but few of them explicitly consider the spatial component in their analysis (Szumik et al. 2002;Echeverry and Morrone 2010). Szumik et al. (2002) and Szumik and Goloboff (2004) proposed and applied an optimality criterion (endemicity analysis; EA) to the delimitation of Ae, which is defined as follows: 'a species could be counted as endemic in an Area A when it satisfies the requirement (evenness rule) that it is present in at least two cells of A, and in each cell C of the area the species is either present, or absent in no more than Q (where 0 < Q < 8) of the cells around C that belong to A' (Szumik and Goloboff 2004). ...
Article
Full-text available
The mammals are the biological group initially analysed by Wallace to define the Neotropical region (NR). Their areas of endemism (Ae) are considered historical patterns, which have been used to describe biogeographic schemes. However, the Ae at regional scale are currently unclear. In the present study, we analyse Ae of mammals at the regional scale and compare them with previous biogeographic schemes of the NR. The Ae of Neotropical terrestrial mammals were identified using the endemicity analysis (software NDM/VNDM). Our results showed that the NR is composed of 10 Ae, supported by 82 endemic taxa (6 families, 29 genera, and 47 species). The Ae showed a NR with multiple boundaries and with a core of higher overlap of the areas of endemism (OAE) from Veracruz and the Pacific coasts of Mexico to the southern limit of Amazonia in Brazil. The NR boundaries vary strikingly with latitude, with substantially more overlapping areas of endemism in the tropical biomes than in the temperate biomes of America. This pattern of OAE is consistent with the higher mammal-species richness zone within the tropical biomes and other biogeographic patterns such as higher productivity and spatial heterogeneity.
... KEY WORDS. -chelonians; monospecific genera; bispecific genera; endemism index; island; archipelagoes; macroecology; conservation Evolutionary theory suggests that mono-and bispecific genera are noteworthy compared with multispecies genera because they either may be evolutionarily older (Ridley 1993) or would represent not-yet-diversified young lineages (Krug et al. 2008). In addition, many of the mono-and bispecific genera are endemic, in some cases confined to islands and archipelagoes (Ridley 1993). ...
... -chelonians; monospecific genera; bispecific genera; endemism index; island; archipelagoes; macroecology; conservation Evolutionary theory suggests that mono-and bispecific genera are noteworthy compared with multispecies genera because they either may be evolutionarily older (Ridley 1993) or would represent not-yet-diversified young lineages (Krug et al. 2008). In addition, many of the mono-and bispecific genera are endemic, in some cases confined to islands and archipelagoes (Ridley 1993). For example, of 83 genera of island endemic rodents (Amori et al. 2008), 57 (68.7%) were mono-and bispecific (Amori et al. 2008(Amori et al. , 2017. ...
... For all analyses, fossil species were excluded. The biogeographical region of each taxon was classified according to Wallace (1894), revised in Rueda et al. (2013). In this regard, we used the term ''Afrotropical'' instead of ''Ethiopian,'' which was used originally by Wallace (1894). ...
Article
Mono- and bispecific genera are noteworthy compared to multispecies genera because they either may be evolutionarily older or would represent not-yet-diversified young lineages, and, being in some cases endemic to specific islands and archipelagoes, they also are of conservation concern. Here, the distribution patterns of the mono- and bispecific genera of freshwater turtles and tortoises and the distribution patterns of island endemics are analyzed, using a database provided by the International Union for Conservation of Nature/Species Survival Commission Tortoise and Freshwater Turtle Specialist Group. The mean number of species per genus varied significantly across biogeographic regions. Twenty-eight genera are monospecific, and 18 are bispecific, accounting for 48.9% of the total genera of tortoises and freshwater turtles worldwide. The Oriental region housed the highest fraction of these genera, followed by the Neotropical and Afrotropical regions. More than 11% of the total number of chelonian species (n = 356) were island endemics, with most species being Neotropical. The majority of the endemic island species occurred in the Galapagos Islands and in Papua New Guinea. The endemism index varied remarkably among the various islands/archipelagoes, with the Philippines being the center of endemism with the minor value of the index and the Galapagos being that with the highest value of the index. Island size was correlated neither with the number of endemic species per island nor with the endemism index of each island, but it was significantly correlated with the total number of species (once the Galapagos were removed from analyses as an outlier). Species belonging to mono- and bispecific genera, as well as island endemic species, were more threatened (as a percentage; in all cases . 65%) than the overall percentage of threatened taxa in freshwater turtles and tortoises worldwide (50.3%).
... Wallace went further to discuss the complex process of dividing the Earth into regions based on the distribution of species and some of the principles he employed in his regionalization scheme. He indicated that regions should follow natural divisions as close as possible but should also be approximately the same size (Rueda et al., 2013). The primary rule for delineating a region was the presence of characteristic genera. ...
... Nevertheless, once these parameters are specified, the process is repeatable, and the results most often make intuitive sense (Fig. 2). Rueda et al. (2013) proposed using adaptive affinity propagation to determine the optimal number of clusters instead of using arbitrary values as the optimal number of clusters. Species turnover indices are most commonly used in cluster analyses, however, there are some concerns when using these indices to differentiate regions. ...
... Phylogenetic trees provide important information about the evolutionary relationships between species, allowing for schemata to be derived fulfilling Wallace's original vision (Holt et al., 2013;Daru et al., 2017). In the past, regions were delineated based on species distributions and the presence (or absence) of characteristic genera (Wallace, 1876;Rueda et al. 2013). Modern developments have led to the delineation of biogeographic regions based on evolutionary history (i.e., phylogenetically important regions), which is centered on the concept of radiation of a single monophyletic clade which promotes high levels of endemism (Kreftz and Jetz, 2013;Holt et al. 2013;Maestri and Duarte, 2020). ...
Chapter
Regionalization is the discipline that groups geographic areas of the world into regions based on predetermined criteria. While these may be abiotic criteria (e.g., climate), in biogeography the term (more specifically referred to as bioregionalization), has essentially come to mean that regions are defined on the basis of distribution patterns in living organisms. Most often this has been done using species of tetrapod vertebrates and vascular plants, with other studies considering other taxa, or shared branch lengths in the phylogenies for these groups. This process was initially performed intuitively, but currently, this is achieved using increasingly sophisticated algorithms. The results for approaches using different methods and looking at different taxa tend to converge towards common global schemata which can be explained using past and present climate, plate tectonics, and the evolution of life on Earth. Preserving distinctive assemblages of living organisms, as illustrated in regionalisation exercises, is increasingly viewed as one important facet of biodiversity conservation.
... But, paradoxically, several biogeographical regionalizations based on the analysis of large databases have been published recently (e.g. Escalante, 2017;Holt et al., 2013;Kreft & Jetz, 2010;Proches ß & Ramdhani, 2012;Ribeiro et al., 2014;Rueda, Rodr ıguez, & Hawkins, 2013). It seems that biogeographical regionalization is back, like a spectre, haunting molecular/parametric biogeographers and others who ignore biogeographical regionalizations. ...
... Proches ß and Ramdhani (2012) considered that genus-level data are preferable to species-or family level data, because species-level taxonomy is often debatable and family level classification may change substantially, so genera are the best choice (see also Rueda et al., 2013). Furthermore, Proches ß and Ramdhani (2012) considered that the number of genera is not so large to raise computational problems and despite differences in lineage age they can be adequate surrogates in cross-taxon analyses. ...
Article
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A biogeographical regionalization is a hierarchical system that categorizes geographical areas in terms of their biotas. I provide a general protocol to undertake biogeographical regionalizations, that consists of seven steps: (1) defining the study area; (2) assembling distributional data; (3) identifying natural areas; (4) discovering area relationships; (5) defining boundaries/transition zones; (6) regionalization and (7) area nomenclature. Natural biogeographical units are useful for people undertaking different types of analyses, like macroecologists, evolutionary biologists, systematists and conservationists. Biogeographical regionalizations may help biogeographers communicate more effectively between themselves and discover opportunities to work on common problems, contributing to the development of a truly integrative biogeography.
... Rooted in the seminal ideas of Alexander von Humboldt, this fascination has promoted a long-term research agenda aiming to delineate biogeographic regions according to their faunas and floras (e.g. [1][2][3]). Besides this, the large-scale eco-evolutionary processes that shape regional biotas are known to influence ecological and evolutionary dynamics at finer scales [4]. ...
... All rights reserved. early and mid-Cenozoic, species distributions would be more influenced by recent events such as Pleistocene glaciations [3]. These recent events could have promoted many additions and subtractions of species to regional faunas through dispersal and diversification processes. ...
Article
Although the description of bioregions dates back to the origin of biogeography, the processes originating their associated species pools have been seldom studied. Ancient historical events are thought to play a fundamental role in configuring bioregions, but the effects of more recent events on these regional biotas are largely unknown. We used a network approach to identify regional and sub-regional faunas of European Carabus beetles and developed a method to explore the relative contribution of dispersal barriers, niche similarities and phylogenetic history on their configuration. We identify a transition zone matching the limit of the ice sheets at the Last Glacial Maximum. While southern species pools are mostly separated by dispersal barriers, in the north species are mainly sorted by their environmental niches. Strikingly, most phylogenetic structuration of Carabus faunas occurred during the Pleistocene. Our results show how extreme recent historical events-such as Pleistocene climate cooling, rather than just deep-time evolutionary processes-can profoundly modify the composition and structure of geographical species pools.
... A large stretch of barren desert, acting as a zoogeographical barrier, lies between them. Boundaries among zoogeographical realms in the Arabian Peninsula is continuously a trending controversial topic (Larsen 1984;Rueda et al. 2013;El-Hawagry and Al Dhafer 2015;Ficetola et al. 2017;El-Hawagry et al. 2019). ...
... These distributional patterns indicate that zoogeographically the area of the Arabian Peninsula is not a homogeneous unit. Our analysis of Lepisiota zoogeographic affinities generally supports Larsen (1980), Larsen (1984), Abdel-Dayem et al. (2019), Cowie (1989), Penati and Vienna (2006), Rueda et al. (2013), Sharaf et al. (2014), Ficetola et al. (2017, Delany (1989), El-Hawagry et al. (2019 arguments that a major zoogeographic discontinuity exists within the region. Despite this, as mentioned above about 38% (8 out of 21 spp.) of the species appear to be endemic to the region. ...
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This study updates and summarizes information on the taxonomy and status of the Arabian Lepisiota fauna. We describe and illustrate the new species Lepisiota elbazisp. nov. from the Dhofar Governorate, Oman based on the worker caste. The new species is closest to the Arabian species, L. arabica Collingwood, 1985 from the southwestern mountains of the Kingdom of Saudi Arabia (KSA) and can be separated by having fewer body hairs (two pairs on the posterior margin of the head, two or three pairs on the promesonotum and, one or two pairs on the first gastral tergite), the longer head, scapes, and propodeal spines, and the shorter mesosoma. We present the first illustrated key to the worker caste of the Arabian species of Lepisiota using stacked digital color images to facilitate species determination. The new species is probably endemic to the Dhofar Governorate and seems rare. An up-to-date synoptic checklist of 21 species representing the Arabian Lepisiota Santschi, 1926 is emended based upon the most recent literature in ant systematics. Five species are excluded from the Arabian Lepisiota fauna, L. arenaria (Arnold, 1920), L. erythraea (Forel, 1910), L. incisa (Forel, 1913), L. sericea (Forel, 1892a), and L. simplex (Forel, 1892) for issues related to previous species misidentification. Lepisiota carbonaria (Emery, 1892) is proposed as a senior synonym of L. depilis (Emery, 1897) syn. nov. The faunal composition of Lepisiota species recorded from the Arabian Peninsula can be divided/delineated into two main groups according to their zoogeographical relationships; (1) Afrotropical (11 species-~52.38%); (2) Palearctic (10 species-~47.62%) elements whereas eight species (~38%) are Arabian endemics.
... It is a transition zone between Sahul and Sundaland. Halmahera and Papua (New Guinea) were the part of Australian zoogeographic region (Wallace 1876;Rueda et al. 2013) which were divided by Holt et al. (2013) into Australian and Oceanian including Moluccas, New Guinea, and Pacific Islands. ...
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Nyctimystes infrafrenatus Günther is a widespread frog species distributed across northern Australia, New Guinea, Bismarck archipelago and in the eastern part of Indonesia archipelago where the species thus encompasses two biogeographic regions, Papua and Wallacea. We evaluated phylogenetic relationships among the populations from Papua and Moluccas based on two mitochondrial loci (16S rRNA and Cytochrome Oxidase Subunit I). Two major subclades are recovered within Nyctimystes infrafrenatus with subclade A being represented by populations from New Guinea and northern Australia and subclade B by Moluccan populations (Halmahera and Tidore). Genetic distances (2.3-4.2 in 16S rRNA) between these subclades suggest they could belong to distinct species. Since New Guinea populations correspond to the nominal species and that Nyctimystes tennuigranulatus (Boettger 1895), currently considered as a junior synonym of Nyctimystes infrafrenatus, is available for the northern Moluccan populations, we proposed to remove this taxon from synonymy. However, samples used in this study come from northern Moluccas, further studies including sample from Ambon is needed to determine taxonomic status of the southern Moluccan population whether they are conspecific with northern Moluccan population or not.
... However, amphibians, birds and mammals have different life-history traits, including metabolism, eco-physiological tolerance, and dispersal abilities. Several analyses have shown that bioregions designed on the basis of different vertebrate clades show a significantly different structure, with clear variability in the number of regions and in the positions of boundaries (e.g., Rueda et al., 2010;Holt et al., 2013;Rueda et al., 2013;Edler et al., 2016; see Supporting Information Figure S1). It is likely that groups with different features (e.g., dispersal, metabolism, or evolutionary history) show divergent responses to biogeographical factors (Lomolino et al., 2010). ...
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Aim Worldwide distribution patterns of living animals are structured in multiple zoogeographical regions, characterized by faunas with homogeneous composition that are separated by sharp boundaries. These zoogeographical regions can differ depending on the considered animal group, probably because they have distinct characteristics such as dispersal, metabolism, or evolutionary history, and thus divergent responses to major biogeographical drivers, such as tectonic movements, abrupt climate transitions and orographic barriers. Here, we tested if the drivers of biogeographical boundaries are different between vertebrate classes with strongly divergent traits and evolutionary history. Location Global. Time period Present. Major taxa studied Amphibians, birds and mammals. Methods We focused on terrestrial biogeographical boundaries, considering multiple potential drivers: spatial heterogeneity of present-day climate, altitudinal variation, long-term tectonic movements and past climate change (temperature). We used spatially explicit regression models and geographically weighted regressions to select and quantify the factors explaining the position of the biogeographical boundaries between vertebrate classes. Results For mammals, tectonic movements, abrupt climatic transitions and orographic barriers jointly determined extant biogeographical boundaries, with tectonic movements being the most important. For birds, abrupt climatic transitions played the strongest role, while the effect of orographic barriers was weak. For amphibians, biogeographical boundaries mostly corresponded to areas with abrupt climatic transitions. The strongest transitions of amphibian faunas occur in areas with abrupt shifts of temperature and precipitation regimes. Main conclusions Our analyses confirmed that different drivers have jointly shaped the global vertebrate biogeographical regions, and highlight that taxa with different features show heterogeneous responses across the globe. Eco-physiological constraints likely increase the importance of spatial heterogeneity of climate, while dispersal limitations magnify the relevance of physical barriers (mountain chains and long-term tectonic instability). Integrating among-taxa heterogeneity into analyses thus provides a more complete view of how different processes determine biodiversity variation across the globe.
... Amblyomma geoemydae occurs within the Oriental region, including its northernmost extent: Japan (Ryukyu islands), southern China, and Taiwan (Yamaguti et al., 1971;Keirans and Durden, 2001;Robbins, 2005;Takahashi et al., 2017). Although Guglielmone et al. (2014) regard these areas as the extreme southeast of the Palearctic region, many biogeographers regard these as the northeast reaches of the Oriental region (Sclater 1896;Newton and Dale 2001;Rueda et al., 2013). To date, A. geoemydae has been reported from the following nations: Cambodia, China (PRC), India, Indonesia, Japan (Ryukyu Islands), Malaysia, Myanmar, Philippines, Singapore, Sri Lanka, Taiwan, Thailand, and Vietnam (Cantor, 1847;Yamaguti et al., 1971; Vijaya, 1983; Keirans and Durden, 2001;Robbins et al., 2006;Kwak, 2018;Petney et al., 2019). ...
Article
Amblyomma geoemydae is an oriental tick species primarily associated with turtles and tortoises but also sometimes reported from other vertebrates, including humans. Although A. geoemydae was first reported from the Philippines more than 70 years ago, it has not been reported since, and no local distributional or host data have been published. Herein, we present the first definitive locality report and host record for A. geoemydae in the Philippines. A complete checklist of the known host species from which A. geoemydae has been collected is also presented.
... Additionally, two transition zones have been recognized: the Mexican transition zone in the overlap between the Nearctic and Neotropical regions, and the South American transition zone between the Neotropical and Andean regions . Some recent quantitative analyses have recovered the Neotropical region as the Sclater-Wallace system Holt et al., 2013a;Rueda et al., 2013); however, they are based only on vertebrate taxa, usually at the species level. The analysis of Procheş and Ramdhani (2012) showed a more restricted Neotropical region, excluding southern South America. ...
Book
Neotropical Biogeography: Regionalization and Evolution presents the most comprehensive single-source treatment of the Neotropical region derived from evolutionary biogeographic studies. The book provides a biogeographic regionalization based on distributional patterns of plant and animal taxa, discusses biotic relationships drawn from track and cladistic biogeographic analyses, and identifies cenocrons (subsets of taxa within biotas identified by their common origin and evolutionary history). It includes maps, area cladograms and vegetation profiles. The aim of this reference is to provide a biogeographic regionalization that can be used by graduate students, researchers and other professionals concerned with understanding and describing distributional patterns of plants and animals in the Neotropical region. It covers the 53 biogeographic provinces of the Neotropical region that are classified into the Antillean, Brazilian and Chacoan subregions, and the Mexican and South American transition zones.
... riborus Förster, 1869 is a moderately species rich genus of the family Ichneumonidae, subfamily Campopleginae with 56 valid species worldwide, including the presently described new species; it is most diverse in the Oriental and Eastern Palaearctic regions (Yu et al. 2012). Regarding Eriborus, the biogeographical scope of this work is the Australian (Australasian) realm, including Australia, New Guinea, New Zealand and the surrounding islands (such as New Caledonia or Vanuatu) eastwards from Wallace's line; Fiji and Samoa Islands are also considered here (Rueda et al. 2013), though sometimes these are assigned to the Oceanic realm (see e.g. Olson et al. 2001, Yu et al. 2012. ...
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Eriborus mirabilis sp. nov. is described from Papua New Guinea, and the first identification key to the Australasian species of Eriborus Förster, 1869 is provided. Nepiesta cruella sp. nov. is described from Jordan, the female of Nepiesta tibialis Horstmann, 1977 is described for the first time, and an updated identification key to all known Nepiesta Förster, 1869 species is given. Additionally, the first records of Eriborus obscuripes Horstmann, 1987 from Romania, Eriborus terebrator Aubert, 1960 from Hungary and Spain, Nepiesta mandibularis (Holmgren, 1860) from Hungary and Romania, Nepiesta rufocincta Strobl, 1904 from Romania, and Nepiesta tibialis Horstmann, 1977 from Turkmenistan are reported.
... For all analyses, fossil species were excluded. The zoogeographical region of each taxon was classified according to Wallace (1894), revised in Rueda et al. (2013). In this regard, we used the term 'Afrotropical' instead of 'Ethiopian', which was used originally by Wallace (1894). ...
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Monospecific and bispecific genera are of particular interest in studies of taxonomic diversity and speciation evolution. Here, the distribution patterns of mono- and bispecific amphibians worldwide are investigated, with some discussion of on the conservation implications also presented. Based on an online database search (available from the American Museum of Natural History, New York), we found that the mean number of mono- and bispecific genera was similar among zoogeographic regions, with a total of 120 mono- (95 Anura, 17 Caudata, 8 Gymnophiona) and 65 bispecific (48 Anura, 10 Caudata, 7 Gymnophiona) genera. Out of 73 known amphibian families worldwide, only 35.6% of them do not contain any mono- or bispecific genera. The frequency of mono- or bispecific genera by family was not significantly different among Anura, Caudata and Gymnophiona. There was a general tendency for the number of mono- and bispecific genera of amphibians to be positively correlated with the total number of genera in that family. In Anura, there was a preponderance of mono-specific genera in Afrotropical and Neotropical regions. Concerning bispecific genera, there was a clear preponderance in the Neotropical region for anurans. There was a positive correlation between the number of threatened genera (according to the IUCN Red List) in both the mono- and bispecific groups and the relative number of species in each taxon, thus showing that taxonomical speciosity clearly influences the frequency of occurrence of mono- and bispecific taxa in each family and order. In this regard, Anura dominated in both the number of worldwide described mono- and bispecific taxa as well as in that of the threatened ones according to IUCN Red List.
... This database is the most comprehensive dataset on Iran's mammals and was built from an extensive review of published and unpublished observations (details in Yusefi et al., 2019). To avoid biases resulting from insufficient or uneven sampling, we limited analyses to grid cells with more than five species or with ≥50% of their area within the country borders (Rueda, Rodríguez, & Hawkins, 2013). ...
Article
Aim: In recent years, novel approaches have been proposed to improve current bi-oregionalization methods, but these have not been thoroughly compared. We assessed the applicability of the recently developed network-based clustering method (Infomap algorithm) in bioregionalization analysis at regional spatial scales and compared the results with commonly used distance-based methods (hierarchical clustering algorithm). We also identified climate regions by using a model-based cluster analysis (Gaussian algorithm). Finally, we quantified the representation of climate regions and bioregions in current protected areas (PAs). Location: Iran. Taxa: Terrestrial mammals. Methods: To define bioregions we used the Infomap algorithm and distance-based clustering methods based on species distribution data (over 14,000 occurrence records for 188 species). The Infomap algorithm was applied using the interactive web application "INFOMAP BIOREGIONS" and the distance-based clustering was based on unweighted pair-group method using arithmetic averages (UPGMA). To identify climate regions we used principal components analysis and a model-based cluster analysis both based on 15 climatic variables as well as a terrain ruggedness index. Results: The Infomap algorithm detected nine biogeographical units: seven biore-gions and two transition zones. The distance-based method suggested five biore-gions. The identified bioregions differed between methods with some consistent spatial patterns across methods. Temperature and precipitation explained 85.8% of the environmental variation. Eight climate regions were identified. In general, climate variation and bioregional patterns are currently poorly represented in PAs (<25% coverage). Main conclusions: The network-based method allowed identifying bioregions at regional scale and was apparently more sensitive than the conventional distance-based method. The detection of transition zones by the Infomap algorithm was an advantage, and stressed the fact that the distribution of Iranian mammalian fauna is complex, especially in the southeastern part where contact areas between several bioregions are found. The identified bioregions (especially the distance-based bioregions) and climate regions tended to match well with previous bioregionalization
... These areas agree with studies that attempted to identify historically stable geographic unities of various taxa (Rueda et al., 2013;Varela et al., 2019). ...
Article
An antitropical pattern is characterized by the occurrence of closely related taxa south and north of the tropics but absent or uncommonly represented closer to the equator, in contrast to most taxa, which tend to have their highest diversity in the tropical regions. We investigate the antitropical distribution of eucerine bees with the aim of contributing to the characterization and understanding of this pattern. All continents except Antarctica and Australia. Eucerine bees (Hymenoptera: Apidae: Eucerinae). We carried out phylogenomic dating under two different clock models and used multiple strategies to vary matrix composition, evaluating the overlapping of divergence times estimated across models using Bhattacharyya coefficients. Lastly, we reconstructed the biogeographic history of eucerine bees using a Bayesian implementation of the DEC model. Eucerinae is estimated to have started diversifying during the Palaeocene, with all its tribes originating during the Palaeocene/Eocene transition in southern South America. At least two range expansions happened into North America before the full closure of the Isthmus of Panama. We show that divergence between closely related groups with disjunct distributions would have happened in periods when the climate favoured the expansion of open habitats and became isolated when the forests were re‐established. We describe the early diversification of eucerine bees, revealing an intimate association with southern South America. Events of range evolution of Eucerinae were likely affected by periods of global cooling and aridification, and palaeoclimatic and vegetational conditions probably have been more relevant to the formation of the antitropical distribution of Eucerinae than the consolidation of the Isthmus of Panama connecting the Americas. We also demonstrate that most uncertainty in divergence time estimation is not due to the amount of molecular data being used, but more likely other factors like fossil calibrations and violations of clock models.
... Almost a century and a half ago, Wallace (1876) divided the world into six zoogeographical regions (Fig. 1a) which, to a great extent, coincided with the boundaries of continental tectonic plates. Wallace's point of view is supported by many modern researchers (Ruesta, 2013;Whittaker et al., 2013). The phytogeographical picture of the world (Takhtadzhan, 1978) also largely coincides with Wallace's views. ...
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An analytical overview is presented of the geographical distribution of two testate amoebae of the genera Hoogenraadia and Planhoogenraadia belonging to the Gondwana-tropical group, in the Oriental biogeographic area and Primorskii krai of the Far East. It is shown that some species of these genera occur on the territory of the Sikhote-Alin Nature Reserve and on the islands of Honshu and Hokkaido (Japan), territories belonging to the southeastern part of the Palearctic. Taking into account the geographical distribution of this group of Protozoa, it is proposed to refine the southeastern border of the Palearctic in the Far East area.
... We obtained range maps for all Palaearctic and Nearctic non-volant terrestrial mammal species from (Schipper et al., 2008) and rasterized them on a c. 100 × 100 km equal-area grid comprising 2,244 cells in the Nearctic and 4,162 cells in the Palaearctic. Nearctic and Palaearctic biogeographic regions were delimited according to the mammalian zoogeographic domains in Rueda, Rodríguez, and Hawkins (2013). We used biogeographic regions rather than smaller domains (i.e. ...
Article
We test the ability of the biotic exchange across the Bering land bridge coupled to niche conservatism to explain current day mammalian diversity gradients. The Holarctic. Mammals. We compared the diversity within clades that participated in the exchange (colonizers), whose ancestors withstood the Beringian cold temperatures, with that within clades that did not participate (sedentaries). We contrasted biogeographical patterns, tested the ability of environmental models to predict species richness of colonizers and sedentaries across continents and, compared richness‐climate relationships between colonizers and sedentaries controlling for phylogenetic effects. We find that assemblages of colonizers are more diverse towards higher latitudes, opposing the traditional latitudinal diversity gradient which is followed by sedentaries. Despite the long passage of time since this major dispersal event, we find that the geographic distribution of colonizers is more strongly correlated with the distributions of other colonizers inhabiting a different continent than to the distribution of sedentary species. Our results highlight the importance of historical migrations and dispersal in configuring present‐day diversity gradients. We also suggest that colonizers may be particularly vulnerable to future climate change because of the predicted disproportionate decrease in climate space in the extra‐tropical realm where they are currently most diverse.
... Weksler et al. 2006). Distribution (biogeographical region; following classification in Wallace [1894] revised in Rueda et al. [2013]) and conservation status (in terms of IUCN Red List categories) were drawn from the IUCN database, available at www.iucnredlist.org (accessed on 15 Aug 2015). ...
Article
Mono-and bispecific genera are of special concern as they represent unique phylogenetic/evolutionary trajectories within larger clades. In addition, as phylogenetically older taxa are supposed to be exposed to higher rarity and extinction risk, mono- and bispecific genera may be intrinsically more prone to extinction risks than multispecies genera, although extinction risks also depend on the ecological and biological strategy of the species. Here, the distribution across biogeographical zones and the threatening levels of two speciose orders of mammals (mono- and bispecific genera of Rodentia and Soricomorpha) are investigated in order to highlight major patterns at the worldwide scale. In Rodentia, 39.7% of the genera (n = 490) were monospecific and 17.9% were bispecific. In Soricomorpha, 44.4% of the total genera (n = 45) were monospecific and 15% were bispecific. There was a positive correlation between the number of monospecific genera and the total number of genera per family. Peaks of mono- and bi-specific genera richness were observed in Neotropical, Oriental and Afrotropical regions in rodents and in the Palearctic region in soricomorphs. Range size was significantly uneven across biogeographic region in rodents (with larger ranges in Nearctic and Oriental regions and smaller ranges in the Australian region), but there was no difference across biogeographic regions in terms of range size in soricomorphs. Most of the mono- and bispecific genera occurred in forest habitat in both taxa. The frequency distribution of the mono- and bispecific genera across IUCN categories did not differ significantly from the expected pattern using the total rodent genera and the multispecies genera.
... It separates the Oriental faunal realm to the west from the Australasian one to the east (e.g. Wallace, 1876;Procheş & Ramdhani, 2012;Holt et al., 2013;Rueda et al., 2013). In an attempt to delineate a boundary between the two, Müller (1846) bisected the island chain with an ecology-based line (see also Sarasin & Sarasin, 1901). ...
Article
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Based on a comprehensive literature survey, we determined the sources of the terrestrial vertebrate species on Christmas Island, asking where they originated relative to Wallace’s Line (the southern end of the divide lies 1100 km to the east, where the Lombok Strait adjoins the eastern Indian Ocean). The two bats, Pipistrellus murrayi and Pteropus natalis, are from the west. Concerning the endemic and ‘resident’ bird species, one is from the west (Collocalia natalis), four are from the east (Accipiter fasciatus, Egretta novaehollandiae, Falco cenchroides and Ninox natalis) and the other 15 are ambiguous or indeterminate. Most of the land-locked species are also from the east: rodents Rattus macleari and Rattus nativitatis, and squamates Cryptoblepharus egeriae, Emoia nativitatis and Lepidodactylus listeria. Additionally, two have westerly origins (Crocidura trichura and Cyrtodactylus sadleiri), one is ambiguous (Emoia atrocostata) and another is unknown (Ramphotyphlops exocoeti). West-directed surface currents that flow across the eastern Indian Ocean towards Christmas Island would have facilitated most of the land-animal colonizations. We therefore suggest that Wallace’s Line be redrawn such that the landmass is placed on the Australasian side of this fundamental biogeographical boundary.
... org/10.1016/j.apgeog.2020.102208. Brown and Bredenkamp, 2018, Casazza et al., 2008, Ficetola et al., 2017, Khedr, 2006, Rueda et al., 2013, Violle et al., 2015, Williams et al., 1999 ...
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Biogeographical studies are considered effective for investigations on macroecology and evolution, as well as for delineating patterns of endemism and identifying the key drivers influencing the historical distribution of species. Clustering techniques, based on environmental datasets and distribution of endemic species, have been largely used in biogeography and for the definition of endemic-rich regions where conservation actions should be implemented. Despite several earlier studies have dealt with the biogeographical territories in Egypt, none of them have provided a replicable method to support such regionalisation. We proposed a two-step procedure for the biogeographical regionalisation of Egypt consisting in (1) the definition of environmental clusters (based on 24 climatic, three topographic, two soil organic carbon content and pH, and four habitat heterogeneity variables) by using the k-nearest neighbours algorithm and (2) the spatial clustering of such clusters according to the distribution of 140 endemic vascular plants. The hierarchical clustering and indicator values analyses, based on the presence-absence matrix of endemic taxa in each cluster were performed to define two cut-off levels of biogeographical sectors and subsectors. A total of six sectors and nine subsectors were identified. Climatic-related variables, elevation and soil organic carbon were the most important determinants for environmental clustering of Egypt. The highest endemic richness was recorded in the Marioutico-Arishian (71 species), Sinaico-Arabian (62 species) and Nilotic (16 species) sectors, and in Sinaic (54 species), Arishian (45 species) and Marioutic (40 species) subsectors. Nonetheless, the most protected sectors were Elbanian and Suezian (62.73 and 29.05%, respectively) while the lowest sectors were Nilotic and Marioutico-Arishian (9.86 and 13.26%, respectively). The already established protected areas in Egypt are not sufficient for conserving the representation of the identified endemic rich plant sectors. This two-step procedure confirms the usefulness of environmental attributes together with the spatial distribution of endemic vascular plants to define the biogeographical units in Egypt. Our regionalisation method could be replicated for other species’ groups, with the ultimate goal of integration all species of interest in a single biogeographical system. Furthermore, the presented regionalisation will help to identify weaknesses in current protection actions and to understand biogeographical processes.
... Originally, the Neotropical region was defined according to the Sclater-Wallace system, to comprise South America and part of Central America, extending northbound as far as central Mexico (Sclater, 1858;Wallace, 1876). This scheme has been widely accepted by different zoogeographers studying vertebrate distribution (Cox, 2001;Morrone, 2014Morrone, , 2015Rueda et al., 2013). Other authors, however, especially those interested in invertebrate biogeography, have excluded from the Neotropical region the Andean area and the southern part of South America, as these regions are historically associated with other Austral areas, namely Australia, New Guinea, New Zealand, Tasmania, and South Africa (reviewed by Morrone, 2014Morrone, , 2015. ...
Article
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Vertebrate predation by invertebrates has been classically underexplored and thus underestimated, despite the fact that many arthropods consume vertebrates. To shed some light on the relevance that spider predation may have upon lizards in the Neotropical and Andean regions, we compiled the available information in the literature on this trophic interaction. We found 50 reports of spiders consuming lizards in these regions, and the 88% of these were from the Neotropical region. Spiders belong to eight families, but Ctenidae and Theraphosidae were the most frequently reported predators. Lizards belong to 12 families, and the most commonly consumed species corresponded to the families Dactyloidae (all Anolis lizards), Gymnophthalmidae and Sphaerodactylidae. Data suggest trophic spider-lizard associations between Ctenidae and Dactyloidae, followed by Theraphosidae and Liolaemidae. The body sizes of the spiders and lizards showed a positive relationship, and spiders were smaller than their prey. We conclude that various spider taxa can be considered lizard predators and they may be ecologically important in the Neotropical and Andean regions. However, spiders of prime predation relevance seem to be those of the Ctenidae and Theraphosidae families.
... It separates the Oriental faunal realm to the west from the Australasian one to the east (e.g. Wallace, 1876;Procheş & Ramdhani, 2012;Holt et al., 2013;Rueda et al., 2013). In an attempt to delineate a boundary between the two, Müller (1846) bisected the island chain with an ecology-based line (see also Sarasin & Sarasin, 1901). ...
Article
Based on a comprehensive literature survey, we determined the sources of the terrestrial vertebrate species on Christmas Island, asking where they originated relative to Wallace’s Line (the southern end of the divide lies 1100 km to the east, where the Lombok Strait adjoins the eastern Indian Ocean). The two bats, Pipistrellus murrayi and Pteropus natalis, are from the west. Concerning the endemic and ‘resident’ bird species, one is from the west (Collocalia natalis), four are from the east (Accipiter fasciatus, Egretta novaehollandiae, Falco cenchroides and Ninox natalis) and the other 15 are ambiguous or indeterminate. Most of the land-locked species are also from the east: rodents Rattus macleari and Rattus nativitatis, and squamates Cryptoblepharus egeriae, Emoia nativitatis and Lepidodactylus listeria. Additionally, two have westerly origins (Crocidura trichura and Cyrtodactylus sadleiri), one is ambiguous (Emoia atrocostata) and another is unknown (Ramphotyphlops exocoeti). West-directed surface currents that flow across the eastern Indian Ocean towards Christmas Island would have facilitated most of the land-animal colonizations. We therefore suggest that Wallace’s Line be redrawn such that the landmass is placed on the Australasian side of this fundamental biogeographical boundary.
... The dividing of the world into 'natural' biodiversity regions, defined as localities with a consistency and distinctiveness of the flora and fauna, has attracted the attention of scientists since the early 19th century [1]. The great islands of western Indonesia (Sumatra, Indonesian Borneo and Java) lie close to the southeastern corner of Asia [2]. ...
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The Tawny coster Acraea terpsicore (Lepidoptera: Nymphalidae) is a non-native species of butterfly that has been recorded in Sumatra since 2009. Summarize recent review confirmed 40 spatial distribution records of A. terpsicore between 2009 to 2020. These records suggest A. terpsicore have widely distributed and colonialized in Sumatra.
... The organisation of the Afrotropical biota in subregions is an ancient theme (Sclater, 1858;Wallace, 1876;Engler, 1879). With the advancement of the informatisation and availability of online databases, the discussion increased and incorporated more taxonomic groups (Cox, 2001;Kreft & Jetz, 2010;Rueda et al. 2013;Ribeiro et al. 2014;Escalante, 2017;Liria et al. 2021). The regionalisation effort under processual and cladistic views allows a more informative biogeographical scenario (Morrone, 2015). ...
Article
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This article explores the distributional data of 2440 occurrence records for 1415 Tipulomorpha (Insecta: Diptera) species to search endemism patterns in the Afrotropical region and to propose a regionalisation scheme. The results were compared to areas of endemism proposed in previous works for other taxa. Areas of endemism were established using NDM/VNDM based on a grid size of 8° and consensus cut-off of 24%. In this study, five subregions were found: Arabian, Atlantic, Indian, Southern Africa and Central Africa. These subregions are divided into 12 dominions: Arabian Peninsula, Atlantic Islands, Central Africa, Central Madagascar, Eastern Africa, Indian Ocean, Northwestern Africa, Southeastern Africa, Southern Africa, Southern Madagascar, Southwestern Africa and Western Africa. There is a general congruence of the dominions with the distribution of the main hydrographic basins and divisions of the forests of Madagascar.
... Distribution patterns of threatened terrestrial vertebrates generally followed the topographical features, mountainous regions show high richness owing to the abundant microhabitats generated by the heterogeneity in climate, rapid elevational changes and varying aspects of slope direction (Spehn and Körner, 2005;Tang et al., 2006). In addition, the Hengduan Mountains also represents the intersection of the Sino-Japanese and Oriental Kingdoms, and historical processes and the divergence of species have also contributed to the high diversity and endemism (Holt et al., 2013;Rueda et al., 2013). Species richness, endemism, and taxonomic distinctiveness are often taken as different dimensions of measuring biodiversity (Myers et al., 2000;Jenkins et al., 2013). ...
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As factors of distribution of Arachnida are outlined paleogeography and paleodistribution, age of groups, barriers, bridges, ability to overcome them, phoresy, dispersal, climate, orography and many other fundamental concepts.
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To improve the phytogeographic characterization of peninsular Italy, we analysed and compared the spatial distribution of woody flora at species and genus levels along an approximate 160 km west–east transect within the central peninsula. Sampling sites were selected using a subjective stratified sampling design. A total of 1710 occurrence data, 138 species, and 74 genera from 153 floristic relevés was used. The congruence in species and genera patterns, supported by Mantel and Procrustes tests, confirmed the validity of the higher‐taxon approach to analyse woody flora patterns. Nonmetric multidimensional scaling and Pearson's chi square test pointed out that elevation and related bioclimatic factors were the most important drivers of woody species composition. Three well defined woody flora types (coastal low‐elevation, inland middle‐elevation, and inland high‐elevation) were identified using partition around medoids clustering; the species‐rich inland middle‐elevation type supports the recently described Transition unit between the Mediterranean and Eurosiberian Regions. Contrary to previous assumptions by other authors, floristic and chorological differences between the Tyrrhenian and the Adriatic subtypes were detected only in the inland high‐elevation type and not in the coastal low‐elevation one. The adopted methodological approach, based on higher‐taxon approach, on clustering and on ordination of floristic relevés, resulted to be valuable to detect woody flora distribution patterns and to delineate the phytogeographic characters of a very heterogeneous and biodiverse area. This article is protected by copyright. All rights reserved.
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The interest in recognizing spatial patterns of species co-distributions has long led biogeographers and macroecologists to classify the world in biogeographic regions. In this chapter, we aimed to identify regions with distinct species pools, thus representing different biogeographic regions with co-occurring species of anurans in South America. Using quantitative and clustering methods, we recognized six anuran biogeographic regions in South America: two regions are predominantly tropical (named as AMAZON and DIAGONAL-AF); two regions are associated to the Andes mountains (named as MID-ANDES and NORTH-/SOUTH-ANDES); and two regions are broadly located south of the Tropic of Capricorn (named as SUB-TROPICAL and TEMP-GRASS). Using regression and variation partitioning analyses, the six distinct biogeographic regions are mainly predicted by differences in climatic gradients among the biogeographic regions (e.g., clusters located in the different tropical, subtropical, and temperate regions). Yet, the combination of rough topography and habitat structure of major biomes was also a good predictor for other biogeographic regions (e.g., the recognition of the different Andean biogeographic regions having different major biomes, such as montane forests and grasslands).
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A foundational paradigm in biological and Earth sciences is that our planet is divided into distinct ecoregions and biomes demarking unique assemblages of species. This notion has profoundly influenced scientific research and environmental policy. Given recent advances in technology and data availability, however, we are now poised to ask whether ecoregions meaningfully delimit biological communities. Using over 200 million observations of plants, animals and fungi we show compelling evidence that ecoregions delineate terrestrial biodiversity patterns. We achieve this by testing two competing hypotheses: the sharp-transition hypothesis, positing that ecoregion borders divide differentiated biotic communities; and the gradual-transition hypothesis, proposing instead that species turnover is continuous and largely independent of ecoregion borders. We find strong support for the sharp-transition hypothesis across all taxa, although adherence to ecoregion boundaries varies across taxa. Although plant and vertebrate species are tightly linked to sharp ecoregion boundaries, arthropods and fungi show weaker affiliations to this set of ecoregion borders. Our results highlight the essential value of ecological data for setting conservation priorities and reinforce the importance of protecting habitats across as many ecoregions as possible. Specifically, we conclude that ecoregion-based conservation planning can guide investments that simultaneously protect species-, community- and ecosystem-level biodiversity, key for securing Earth’s life support systems into the future. © 2018, The Author(s), under exclusive licence to Springer Nature Limited.
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The analysis of regional scale patterns of diversity allows insights into the processes that have shaped modern biodiversity at the macro‐scale. Previous analyses studying biogeographic regionalisation across different high‐level taxa have shown similar trends at a global scale. However, incorporating phylogenetic methods when comparing biogeographic regionalisation between subgroups facilitates identification of mechanisms leading to the biogeographic distribution of specific taxa. We analysed the spatial trends of phylogenetic diversity and phylogenetic endemism of 325 species of New World bats, using updated range maps of the modern distributions. These analyses showed phylogeographic signals that reflect the different evolutionary histories of these families. Zoogeographical zones were detected based on range‐weighted phylogenetic turnover. Values of high phylogenetic diversity and endemism were distributed differently across families, suggesting niche conservatism, but a general latitudinal trend of diversity was evident across taxa. Overall, two main bioregions were shared across New World bat taxa (Nearctic and Neotropical), with two additional subregions (Andean and La Platan). We found strong support for an additional transitional zone in the Pacific coast of South America for Emballonuridae and Molossidae. Differences in regionalisation across families indicate that niche conservatism, in situ diversification and dispersal ability are major drivers for the regionalisation of New World bats, within a dual‐centre of diversification scenario. We also found strong inter‐familial support for an independent Caribbean biogeographic region. This article is protected by copyright. All rights reserved.
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Zoogeographical regions, or zooregions, are areas of the Earth defined by species pools that reflect ecological, historical and evolutionary processes acting over millions of years. Consequently, researchers have assumed that zooregions are robust and unlikely to change on a human timescale. However, the increasing number of human‐mediated introductions and extinctions can challenge this assumption. By delineating zooregions with a network‐based algorithm, here we show that introductions and extinctions are altering the zooregions we know today. Introductions are homogenising the Eurasian and African mammal zooregions and also triggering less intuitive effects in birds and amphibians, such as dividing and redefining zooregions representing the Old and New World. Furthermore, these Old and New World amphibian zooregions are no longer detected when considering introductions plus extinctions of the most threatened species. Our findings highlight the profound and far‐reaching impact of human activity and call for identifying and protecting the uniqueness of biotic assemblages.
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This study aimed to establish if the Lower R ıo de la Plata Basin (LRPB) wetlands can be considered a biogeographic unit. The species of this area were compiled and segregated according to the habitat, selecting only 87 endemic taxa restricted to the LRPB and linked to wetlands. Distributional data of species obtained from the literature, web databases, biological collections, and field trips were georeferenced. The areas of endemism were established as those areas where the distribution of two or more taxa overlaps in groups of rivers' sections with geographic continuity and were tested with a cluster analysis. This congruence is due to ecological, geomorphological, and historical factors. Four areas of endemism were found: a broad area that comprises the whole study area (Riverine district), which is divided into three nested smaller areas (Paraguay-Paran a Flooding Valleys, Uruguay Basin, and Paran a Delta subdistricts). Then, we analysed 170 taxa distributions to evaluate the relationship between the study area and the neighbouring biogeographic units. According to the results, the study area belongs to the Paran a biogeographic province. Some areas of endemism are hidden inside broader areas and are hardly detected with the currently used biogeographic grid-methods. We propose to combine the information about ecological requirements of each taxon with its georeferenced records to estimate their areas of distribution as a primary step for searching areas of endemism in intracontinental studies.
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A new method, multivariate similarity clustering analysis (MSCA) method, was established for biogeographical distribution analyzing. General similarity formula (GSF), the core of MSCA method, can be used to calculate the similarity coefficients between two and among any ≥ three geographical units. Taking the global insects as example, we introduced the steps to use of GSF and consequent clustering processes of this method in details. Firstly, geographical distributions of certain taxa (e.g., Insecta) were categorized into basic geographical units (BGUs); Secondly, similarity coefficients between two and among n BGUs were calculated using GSF; Thirdly, hierarchical clustering was conducted according to values of similarity coefficients (from high to low), then a clustering diagram was generated; Finally, A framework of biogeographical division map was established for the target taxa (e.g, Insecta). We concluded that the MSCA method was efficiently applied in analyzing the biogeographical distribution of given biological taxa; the geographical regions regarding global insects were categorized into 7 Realms with 20 sub‐Realms based on the results of MSCA method. This article is protected by copyright. All rights reserved
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Chigger mites (Acariformes: Trombiculidae) of Southeast Asia, including Myanmar, Malaysia, Brunei, Singapore, Indonesia, Thailand, Cambodia, Vietnam, and the Philippines have been revised based on reference data and examination of type materials in European collections of chiggers. For 450 species of 49 genera synonymy, collection data on types, lists of known host species and lists of countries are given. The lists of hosts include in total 649 valid species and subspecies of mammals, birds, reptiles, amphibians, and arthropods. Two new synonyms were established: Doloisia (Doloisia) Oudemans, 1910 (= Doloisia (Trisetoisia) Vercammen-Grandjean, 1968, syn. nov.) and Gahrliepia lui Chen and Hsu, 1955 (= Gahrliepia (Gateria) octosetosa Chen, Hsu and Wang, 1956, syn. nov.). Twenty-seven new combinations were proposed: Walchia (Ripiaspichia) biliranensis (Brown, 1997), comb. nov., Walchia (Ripiaspichia) huberti (Upham and Nadchatram, 1968), comb. nov., Walchia (Ripiaspichia) parmulaseta (Brown, 1997), comb. nov., and Walchia (Ripiaspichia) serrata (Brown and Goff, 1988), comb. nov., transferred from Gahrliepia Oudemans, 1912; Farrellioides consuetum (Womersley, 1952), comb. nov. (originally in Trombicula Berlese, 1905), Farrellioides nakatae (Nadchatram and Traub, 1964), comb. nov. (originally in Leptotrombidium Nagayo, Miyagawa, Mitamura and Imamura, 1916), and Farrellioides striatum (Nadchatram and Traub, 1964), comb. nov. (originally in Leptotrombidium), transferred from Euschoengastia Ewing, 1938; Guntheria (Phyllacarus) bushlandi (Philip, 1947), comb. nov. (originally in Ascoschoengastia Ewing, 1946), transferred from Guntherana Womersley and Heaslip, 1943 (syn. of Guntheria Womersley, 1939); Kayella masta (Traub and Sundermeyer, 1950), comb. nov. (originally in Ascoschoengastia), transferred from Cordiseta Hoffmann, 1954; Neoschoengastia stekolnikovi (Kalúz, 2016), comb. nov., transferred from Hypogastia Vercammen-Grandjean, 1967; Susa chiropteraphilus (Brown, 1997), comb. nov., Susa masawanensis (Brown, 1998), comb. nov., and Susa palawanensis (Brown and Goff, 1988), comb. nov., transferred from Cheladonta Lipovsky, Crossley and Loomis, 1955; Ericotrombidium cosmetopode (Vercammen-Grandjean and Langston, 1971), comb. nov., transferred from Leptotrombidium; Eutrombicula gigarara (Brown, 1997), comb. nov., transferred from Siseca Audy, 1956; Microtrombicula eltoni (Audy, 1956), comb. nov., transferred from Eltonella Audy, 1956; Trombiculindus alethrix (Traub and Nadchatram, 1967), comb. nov., Trombiculindus cuteanum (Vercammen-Grandjean and Langston, 1976), comb. nov., Trombiculindus frondosum (Traub and Nadchatram, 1967), comb. nov., Trombiculindus hastatum (Gater, 1932), comb. nov., Trombiculindus lepismatum (Traub and Nadchatram, 1967), comb. nov., Trombiculindus limi (Traub and Nadchatram, 1967), comb. nov., Trombiculindus maxwelli (Traub and Nadchatram, 1967), comb. nov., Trombiculindus roseannleilaniae (Brown, 1992), comb. nov., Trombiculindus sarisatum (Traub and Nadchatram, 1967), comb. nov., Trombiculindus vanpeeneni (Hadi and Carney, 1977), comb. nov., and Trombiculindus yooni (Traub and Nadchatram, 1967), comb. nov., transferred from Leptotrombidium.
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Identifying structure underlying high-dimensional data is a common challenge across scientific disciplines. We revisit correspondence analysis (CA), a classical method revealing such structures, from a network perspective. We present the poorly-known equivalence of CA to spectral clustering and graph-embedding techniques. We point out a number of complementary interpretations of CA results, other than its traditional interpretation as an ordination technique. These interpretations relate to the structure of the underlying networks. We then discuss an empirical example drawn from ecology, where we apply CA to the global distribution of Carnivora species to show how both the clustering and ordination interpretation can be used to find gradients in clustered data. In the second empirical example, we revisit the economic complexity index as an application of correspondence analysis, and use the different interpretations of the method to shed new light on the empirical results within this literature.
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Areas of endemism characterize geographical regions by their unique biotas, providing the basis for studies on the ecological and historical drivers of these biologically distinct units. Tribe Bignonieae (Bignoniaceae) are a highly diverse clade of lianas distributed throughout the Neotropics, representing an excellent model for studying the drivers of species diversity and distribution patterns in this region. We used a dataset representing 98% of the diversity of Bignonieae and 21 170 unique locality records to perform an analysis of endemicity using NDM/VNDM. We recovered areas of endemism distributed across the Neotropics, including a higher number of areas at coarser spatial scales. Although overlapping and nested patterns of endemism were common and the spatial congruence with the individual units of previous regionalization schemes was low, the patterns of endemism recovered were in general agreement with those documented for other taxa. Our findings are generally consistent with key Neotropical biogeographical hypotheses. These results highlight the importance of studying detailed distribution patterns of selected taxa for an improved understanding of Neotropical biogeography.
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The distribution of living organisms on Earth is spatially structured. Early biogeographers identified the existence of multiple zoogeographical regions, characterized by faunas with homogeneous composition that are separated by biogeographical boundaries. Yet, no study has deciphered the factors shaping the distributions of terrestrial biogeographical boundaries at the global scale. Here, using spatial regression analyses, we show that tectonic movements, sharp changes in climatic conditions and orographic barriers determine extant biogeographical boundaries. These factors lead to abrupt zoogeographical transitions when they act in concert, but their prominence varies across the globe. Clear differences exist among boundaries representing profound or shallow dissimilarities between faunas. Boundaries separating zoogeographical regions with limited divergence occur in areas with abrupt climatic transitions. In contrast, plate tectonics determine the separation between deeply divergent biogeographical realms, particularly in the Old World. Our study reveals the multiple drivers that have shaped the biogeographical regions of the world.
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Philosophies underlying past classifications of world faunal regions have fostered the view that such units can have little more than heuristic value. Contributing to this feeling is the difficulty attending assessment of rank relationships among the regional elements of a particular system. In the present study, logical constraints upon the meaning of a satisfactorily efficient hierarchical faunal classification are identified and operationalized into a system of world mammal faunal regions via a iterative procedure involving multidimensional scaling. The result is a classification consisting of four regions and ten subregions, each of the latter of which is as unique as it can be with respect to all other subregions while still contributing to logical hierarchical order relationships within the system. The classification is compared to the Sclater-Wallace system and is shown to be both more efficient and more internally consistent. Summary data and statistics pertaining to the new classification are presented and briefly discussed.
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Affinity propagation clustering (AP) has two limitations: it is hard to know what value of parameter 'preference' can yield an optimal clustering solution, and oscillations cannot be eliminated automatically if occur. The adaptive AP method is proposed to overcome these limitations, including adaptive scanning of preferences to search space of the number of clusters for finding the optimal clustering solution, adaptive adjustment of damping factors to eliminate oscillations, and adaptive escaping from oscillations when the damping adjustment technique fails. Experimental results on simulated and real data sets show that the adaptive AP is effective and can outperform AP in quality of clustering results.
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Kreft and Jetz’s critique of our recent update of Wallace’s zoogeographical regions disregards the extensive sensitivity analyses we undertook, which demonstrate the robustness of our results to the choice of phylogenetic data and clustering algorithm. Their suggested distinction between “transition zones” and biogeographic regions is worthy of further investigation but is thus far unsubstantiated.
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Holt et al. (Report, 4 January 2013, p. 74) propose substantial modifications of Wallace’s long-standing zoogeographic regions based on clustering of a pairwise similarity matrix of vertebrate assemblages. We worry about their compromised use of phylogenies and show that a fundamental point of their analysis—i.e., the delineation of new realms—is only weakly supported by their results and conceptually flawed.
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AbstractA recent Guest Editorial by Parenti & Ebach (2013, Journal of Biogeography, 40, 813–820) disagrees with the methods or interpretations in two of our recent papers. In addition, the authors open a debate on biogeographical concepts, and present an alternative philosophy for biogeographical research in the context of their recently described biogeographical subregion called ‘Pandora’. We disagree with their approach and conclusions, and comment on several issues related to our differing conceptual approaches for biogeographical research; namely, our use of molecular phylogenetic analyses, including time estimates; and Parenti & Ebach's reliance on taxon/general area cladograms. Finally, we re‐examine their ‘tests’ supporting the existence of ‘Pandora’.
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Modern attempts to produce biogeographic maps focus on the distribution of species, and the maps are typically drawn without phylogenetic considerations. Here, we generate a global map of zoogeographic regions by combining data on the distributions and phylogenetic relationships of 21,037 species of amphibians, birds, and mammals. We identify 20 distinct zoogeographic regions, which are grouped into 11 larger realms. We document the lack of support for several regions previously defined based on distributional data and show that spatial turnover in the phylogenetic composition of vertebrate assemblages is higher in the Southern than in the Northern Hemisphere. We further show that the integration of phylogenetic information provides valuable insight on historical relationships among regions, permitting the identification of evolutionarily unique regions of the world.
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The world's zoogeographical regions were historically defined on an intuitive basis, with no or a limited amount of analytical testing. Here, we aimed (a) to compare analytically defined global zoogeographical clusters for the herpetofauna, birds, mammals, and all these groups taken together (tetrapod vertebrates); (b) to use commonalities among these groups to propose an updated global zoogeographical regionalization; and (c) to describe the resulting regions in terms of vertebrate diversity and characteristic taxa. The clusters were remarkably uniform across taxa and similar to previous intuitively defined regions. Eleven vertebrate-rich (Nearctic, Caribbean, Neotropical, Andean, Palearctic, Afrotropical, Madagascan, Indo-Malaysian, Wallacean, New Guinean, Australian) and three vertebrate-poor (Arctic, Antarctic, Polynesian) zoogeographical regions were derived; the Neotropical,- Afrotropical, and Australian had the highest numbers of characteristic tetrapod genera. This updated regionalization provides analytically accurate divisions of the world, relevant to conservation, biogeographical research, and geography education.
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The use of analytical techniques to delineate biogeographical regions is becoming increasingly popular. One recent example, Heikinheimo et al. (Journal of Biogeography, 2007, 34, 1053–1064), applied the k-means clustering algorithm to define the biogeography of the European land mammal fauna. However, they used the Euclidean distance measure to cluster grid cells described by species-occurrence data, which is inappropriate. The Euclidian distance yields misleading results when applied to species-occurrence data because of the double-zero problem and the species-abundance paradox. We repeat their analysis using the Hellinger distance, a measure appropriate for species-occurrence data and which has been shown to outperform other such measures. Our results differ substantially from those presented by Heikinheimo et al. We argue that the rigorous application of appropriate statistical techniques is of crucial concern within conservation biogeography.
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Aim Although sharing many similarities in their vegetation types, South America and Africa harbour very dissimilar recent mammal faunas, not only taxonomically but also in terms of several faunistic patterns. However late Pleistocene and mid-Holocene faunas, albeit taxonomically distinct, presented many convergent attributes. Here we propose that the effects of the Holocene climatic change on vegetation physiognomy has played a crucial role in shaping the extant mammalian faunistic patterns. Location South America and Africa from the late Pleistocene to the present. Methods Data presented here have been compiled from many distinct sources, including palaeontological and neontological mammalian studies, palaeoclimatology, palynology, and publications on vegetation ecology. Data on Pleistocene, Holocene and extant mammal faunas of South America and Africa allowed us to establish a number of similar and dissimilar faunistic patterns between the two continents across time. We then considered what changes in vegetation physiognomy would have occurred under the late Pleistocene last glacial maximum (LGM) and the Holocene climatic optimum (HCO) climatic regimes. We have ordained these proposed vegetation changes along rough physiognomic seral stages according to assumptions based on current botanical research. Finally, we have associated our hypothesized vegetation changes in South America and Africa with mammalian faunistic patterns, establishing a putative causal relationship between them. Results The extant mammal faunas of South America and Africa differ widely in taxonomical composition; the number of medium and large species they possess; behavioural and ecological characteristics related to herbivore herding, migration and predation; and biogeographical patterns. All such distinctions are mostly related to the open formation faunas, and have been completely established around the mid-Holocene. Considering that the mid-Holocene was a time of greater humidity than the late Pleistocene, vegetation cover in South America and Africa would have been dominated by forest or closed vegetation landscapes, at least for most of their lower altitude tropical regions. We attribute the loss of larger-sized mammal lineages in South America to the decrease of open vegetation area, and their survival in Africa to the existence of vast savannas in formerly steppic or desertic areas in subtropical Africa, north and south of the equator. Alternative explanations, mostly dealing with the disappearance of South American megamammals, are then reviewed and criticized. Main conclusions The reduction of open formation areas during the HCO in South America and Africa explains most of the present distinct faunistic patterns between the two continents. While South America would have lost most of its open formations within the 30° latitudinal belt, Africa would have kept large areas suitable to the open formation mammalian fauna in areas presently occupied by desert and semi-arid vegetation. Thus, the same general climatic events that affected South America in the late Pleistocene and Holocene also affected Africa, leading to our present day faunistic dissimilarities by maintaining the African mammalian communities almost unchanged while dramatically altering those of South America.
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Aim Biogeographical regionalizations, such as zoogeographical regions, floristic kingdoms or ecoregions, represent categorizations central to many basic and applied questions in biogeography, ecology, evolution and conservation. Traditionally established by experts based on qualitative evidence, the lack of transparency and quantitative support has set constraints on their utility. The recent availability of global species range maps, novel multivariate techniques and enhanced computational power now enable a quantitative scrutiny and extension of biogeographical regionalizations that will facilitate new and more rigorous uses. In this paper we develop and illustrate a methodological roadmap for species-level biogeographical regionalizations at the global scale and apply it to mammals.
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This paper examines how to obtain species biplots in unconstrained or constrained ordination without resorting to the Euclidean distance [used in principal-component analysis (PCA) and redundancy analysis (RDA)] or the chi-square distance [preserved in correspondence analysis (CA) and canonical correspondence analysis (CCA)] which are not always appropriate for the analysis of community composition data. To achieve this goal, transformations are proposed for species data tables. They allow ecologists to use ordination methods such as PCA and RDA, which are Euclidean-based, for the analysis of community data, while circumventing the problems associated with the Euclidean distance, and avoiding CA and CCA which present problems of their own in some cases. This allows the use of the original (transformed) species data in RDA carried out to test for relationships with explanatory variables (i.e. environmental variables, or factors of a multifactorial analysis-of-variance model); ecologists can then draw biplots displaying the relationships of the species to the explanatory variables. Another application allows the use of species data in other methods of multivariate data analysis which optimize a least-squares loss function; an example is K-means partitioning.
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The Sahara and Sahel regions of northern Africa have complex environmental histories punctuated by sudden and dramatic regime shifts in climate and ecological conditions. Here we review the current understanding of the causes and consequences of two environmental regime shifts in the Sahara and Sahel. The first regime shift is the sudden transition from vegetated to desert conditions in the Sahara about 5500 years ago. Geologic data show that wet environmental conditions in this region—giving rise to extensive vegetation, lakes, and wetlands—came to an abrupt end about 5500 years ago. Explanations for climatic changes in northern Africa during the Holocene have suggested that millennial-scale changes in the Earths orbit could have caused the wet conditions that prevailed in the early Holocene and the dry conditions prevalent today. However, the orbital hypothesis, by itself, does not explain the sudden regime shift 5500 years ago. Several modeling studies have proposed that strong, nonlinear feedbacks between vegetation and the atmosphere could amplify the effects of orbital variations and create two alternative stable states (or regimes) in the climate and ecosystems of the Sahara: a green Sahara and a desert Sahara. A recent coupled atmosphere-ocean-land model confirmed that there was a sudden shift from the green Sahara to the desert Sahara regime approximately 5500 years ago. The second regime shift is the onset of a major 30-year drought over the Sahel around 1969. Several lines of evidence have suggested that the interactions between atmosphere and vegetation act to reinforce either a wet Sahel or a dry Sahel climatic regime, which may persist for decades at a time. Recent modeling studies have indicated that the shift from a wet Sahel to a dry Sahel regime was caused by strong feedbacks between the climate and vegetation cover and may have been triggered by slow changes in either land degradation or sea-surface temperatures. Taken together, we conclude that the existence of alternative stable states (or regimes) in the climate and ecosystems of the Sahara and Sahel may be the result of strong, nonlinear interactions between vegetation and the atmosphere. Although the shifts between these regimes occur rapidly, they are made possible by slow, subtle changes in underlying environmental conditions, including slow changes in incoming solar radiation, sea-surface temperatures, or the degree of land degradation.
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Nature is the international weekly journal of science: a magazine style journal that publishes full-length research papers in all disciplines of science, as well as News and Views, reviews, news, features, commentaries, web focuses and more, covering all branches of science and how science impacts upon all aspects of society and life.
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Ecology, evolution, and historical events all contribute to biogeographic patterns, but studies that integrate them are scarce. Here we focus on how biotic exchanges of mammals during the Late Cenozoic have contributed to current geographic body size patterns. We explore differences in the environmental correlates and phylogenetic patterning of body size between groups of mammals participating and not participating in past biotic exchanges. Both the association of body size with environmental predictors and its phylogenetic signal were stronger for groups that immigrated into North or South America than for indigenous groups. This pattern, which held when extinct clades were included in the analyses, can be interpreted on the basis of the length of time that clades have had to diversify and occupy niche space. Moreover, we identify a role for historical events, such as Cenozoic migrations, in configuring contemporary mammal body size patterns and illustrate where these influences have been strongest for New World mammals.
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Aim To determine if it is possible to generate analytically derived regionalizations for multiple groups of European plants and animals and to explore potential influences on the regions for each taxonomic group. Location Europe. Methods We subjected range maps of trees, butterflies, reptiles, amphibians, birds and mammals to k-means clustering followed by v-fold cross-validation to determine the pattern and number of regions (clusters). We then used the mean range sizes of species in each group as a correlate of the number of regions obtained for each taxon, and climate and species richness gradients as correlates of the spatial arrangement of the group-specific regions. We also included the pattern of tree clusters as a predictor of animal clusters in order to test the ‘habitat templet’ concept as an explanation of animal distribution patterns. Results Spatially coherent clusters were found for all groups. The number of regions ranged from three to eight and was strongly associated with the mean range sizes of the species in each taxon. The cluster patterns of all groups were associated with various combinations of climate, underlying species richness gradients and, in the case of animals, the arrangement of tree clusters, although the rankings of the correlates differed among groups. In four of five groups the tree pattern was the strongest single predictor of the animal cluster patterns. Main conclusions Despite a long history of human disturbance and habitat modification, the European biota retains a discernable biogeographic structure. The primary driver appears to be aspects of climate related to water–energy balance, which also influence richness gradients. For many animals, the underlying habitat structure, as measured by tree distributions, appears to have a strong influence on their biogeographic structure, highlighting the need to preserve natural forest formations if we want to preserve the historical signal found in geographic distributions.
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Aim Nearly 150 years ago, T. H. Huxley modified Wallace’s Line, including the island of Palawan as a component of the Asian biogeographic realm and separating it from the oceanic Philippines. Although Huxley recognized some characteristics of a transition between the regions, Palawan has since been regarded primarily as a peripheral component of the Sunda Shelf. However, several recent phylogenetic studies of Southeast Asian lineages document populations on Palawan to be closely related to taxa from the oceanic Philippines, apparently contradicting the biogeographic association of Palawan with the Sunda Shelf. In the light of recent evidence, we evaluate taxonomic and phylogenetic data in an attempt to identify the origin(s) of Palawan’s terrestrial vertebrate fauna.
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There is general agreement among scientists that biodiversity is under assault on a global basis and that species are being lost at a greatly enhanced rate. This article examines the role played by biogeographical science in the emergence of conservation guidance and makes the case for the recognition of Conservation Biogeography as a key subfield of conservation biology delimited as: the application of biogeographical principles, theories, and analyses, being those concerned with the distributional dynamics of taxa individually and collectively, to problems concerning the conservation of biodiversity. Conservation biogeography thus encompasses both a substantial body of theory and analysis, and some of the most prominent planning frameworks used in conservation. Considerable advances in conservation guidelines have been made over the last few decades by applying biogeographical methods and principles. Herein we provide a critical review focussed on the sensitivity to assumptions inherent in the applications we examine. In particular, we focus on four inter-related factors: (i) scale dependency (both spatial and temporal); (ii) inadequacies in taxonomic and distributional data (the so-called Linnean and Wallacean shortfalls); (iii) effects of model structure and parameterisation; and (iv) inadequacies of theory. These generic problems are illustrated by reference to studies ranging from the application of historical biogeography, through island biogeography, and complementarity analyses to bioclimatic envelope modelling. There is a great deal of uncertainty inherent in predictive analyses in conservation biogeography and this area in particular presents considerable challenges.
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Aim  To produce a spatial clustering of Europe on the basis of species occurrence data for the land mammal fauna.Location  Europe defined by the following boundaries: 11°W, 32°E, 71°N, 35°N.Methods  Presence/absence records of mammal species collected by the Societas Europaea Mammalogica with a resolution of 50 × 50 km were used in the analysis. After pre-processing, the data provide information on 124 species in 2183 grid cells. The data were clustered using the k-means and probabilistic expectation maximization (EM) clustering algorithms. The resulting geographical pattern of clusters was compared against climate variables and against an environmental stratification of Europe based on climate, geomorphology and soil characteristics (EnS).Results  The mammalian presence/absence data divide naturally into clusters, which are highly connected spatially and most strongly determined by the small mammals with the highest grid cell incidence. The clusters reflect major physiographic and environmental features and differ significantly in the values of basic climate variables. The geographical pattern is a fair match for the EnS stratification and is robust between non-overlapping subsets of the data, such as trophic groups.Main conclusions  The pattern of clusters is regarded as reflecting the spatial expression of biologically distinct, metacommunity-like entities influenced by deterministic forces ultimately related to the physical environment. Small mammals give the most spatially coherent clusters of any subgroup, while large mammals show stronger relationships to climate variables. The spatial pattern is mainly due to small mammals with high grid cell incidence and is robust to noise from other subsets. The results support the use of spatially resolved environmental reconstructions based on fossil mammal data, especially when based on species with the highest incidence.