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

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

Historiographical approaches to biogeography: a critical review.

We performed a critical review of the historiographical studies on biogeography. We began with the pioneering works of Augustin and Alphonse de Candolle. Then, we analyzed the historical accounts of biogeography developed by (1) Martin Fichman and his history on the extensionism-permanentism debate; (2) Gareth Nelson and his critique of the Neo-Darwinian historiography of biogeography; (3) Ernst Mayr, with his dispersalist viewpoint; (4) Alan Richardson, who wrote a microhistory on the biogeographic model constructed by Darwin; (5) Michael Paul Kinch and the ideas discussed in the 19th century about the geographical distribution of living beings; (6) Janet Browne, who highlighted the importance of the pre-Darwinian naturalists; (7) Peter Bowler, who focused mainly on the influence of paleontology on biogeography; (8) James Larson, who looked into the practices of the naturalists of Northern Europe in the late 18th century; and (9) Malte Ebach, who like Larson, was more interested in analysing the practices rather than the ideas of naturalists who studied the geographical distribution of organisms. Finally, these works are compared with each other. There has not been a dominant paradigm in the construction of historical narratives of biogeography; however, they provide a useful context for understanding problems of biogeography that continue to be debated to this day.

Biogeographic regionalization of the Neotropical region: New map and shapefile.

We provide a map and shapefile of the 57 biogeographic provinces of the Neotropical region. Recognition of these provinces is based on their endemic species, but their delimitation on the map is based on ecoregions combining climatic, geological, and biotic criteria. These provinces belong to the Antillean, Brazilian and Chacoan subregions, and the Mexican and South American transition zones. We provide a vector file of the biogeographical regionalization by converting the map into a polygon shapefile and a raster file with all provinces.

Matthew's (1915) climate and evolution, the "New York School of Biogeography", and the rise and fall of "Holarcticism".

Climate and evolution (Matthew, 1915) represents an important contribution to evolutionary biogeography, that influenced several authors, notably Karl P. Schmidt, George S. Myers, George G. Simpson, Philip J. Darlington, Ernst Mayr, Thomas Barbour, John C. Poynton, Allen Keast, Léon Croizat, Robin Craw, Michael Heads, and Osvaldo A. Reig. Authors belonging to the "New York School of Zoogeography" -a research community including Matthew, Schmidt, Myers and Simpson- accepted Matthew's "Holarcticism" (north temperate centers of origin) and the permanence of ocean basins and continents, whereas others, especially panbiogeographers and cladistic biogeographers, were extremely critical and reacted against these ideas. "Holarcticism" has been falsified and rejected by dispersalists and the "New York School of Zoogeography" disappeared in the 1970s. Matthew, however, continues being identified by panbiogeographers and cladistic biogeographers as a key representative of classic dispersalism, helping provide some cohesion to their research communities.

Phylogenetic analysis of the family Megalopodidae (Coleoptera: Chrysomeloidea): better taxon-sampling facilitates detection of new relationships and new taxa.

The systematics of Megalopodidae is not adequately known, in spite of it being a relatively small group of phytophagous beetles. The first phylogenetic analysis of Megalopodidae with a comprehensive generic representation (25 genera of 30 described, 10 subgenera and 77 species) is undertaken. A parsimony analysis under equal and implied weights was carried out based on 147 adult and larval morphological characters. Subfamilies Palophaginae and Zeugophorinae were recovered as monophyletic, by contrast with Megalopodinae, which proved to be paraphyletic. Atelederinae are proposed as a new subfamily. Also, three tribes and three subtribes within Megalopodinae are proposed: Leucasteini trib.n., Sphondyliini trib.n. and Megalopodini, the latter including Macrolophina subtrib.n., Temnaspidina subtrib.n. and Megalopodina. The genera Macrolopha, Kuilua, Poecilomorpha, Temnaspis, Antonaria, Agathomerus, Megalopus and Bothromegalopus were recovered as non-monophyletic. New delimitations of the polyphyletic genera Poecilomorpha and Macrolopha are proposed, Clythraxeloma is resurrected, and the subgenera of Agathomerus are suppressed. The following new combinations are proposed: Kuilua apicata (Fairmaire), K. nyassae (Jacoby), Poecilomorpha cribricollis (Pic), P. minuta (Pic), Clythraxeloma assamensis (Jacoby), C. bipartita (Lacordaeri), C. discolineata (Pic), C. downesii (Baly), C. gerstaeckeri (Westwood), C. laosensis (Pic), C. maculata (Pic), C. mouhoti (Baly), C. nigrocyanea (Motschulsky), C. pretiosa (Reineck), Temnaspis tricoloripes (Pic) and Barticaria faciatus (Dalman). Clythraxeloma cyanipennis Kraatz is a restored combination. Distribution patterns of Megalopodidae largely conform to the breakup of Gondwanaland, with its main clades having particular distributions: Andean-Australian (Palophaginae), Ethiopian (Leucasteini, Sphondyliini, and Macrolophina), Neotropical (Ateledrinae and Megalopodina) and Ethiopian-Oriental-Palaearctic (Temnaspidina the result of a secondary expansion. Zeugophorinae present a worldwide distribution, except for the Neotropical and Andean regions, which may be the result of geodispersal. The findings of the present study also shed light on groups with taxonomic issues, where phylogenetic analyses are strongly needed.

Croizat's dangerous ideas: practices, prejudices, and politics in contemporary biogeography.

The biogeographic contributions of Léon Croizat (1894-1982) and the conflictive relationships with his intellectual descendants and critics are analysed. Croizat's panbiogeography assumed that vicariance is the most important biogeographic process and that dispersal does not contribute to biogeographic patterns. Dispersalist biogeographers criticized or avoided mentioning panbiogeography, especially in the context of the "hardening" of the Modern Synthesis. Researchers at the American Museum of Natural History associated panbiogeography with Hennig's phylogenetic systematics, creating cladistic biogeography. On the other hand, a group of New Zealand biologists formalized Croizat's original concepts and soon began arguing with cladistic biogeographers over the relative merits of their approaches. In Latin America, panbiogeography and cladistic biogeography were incorporated as parts of an integrative approach. A recent development, molecular panbiogeography, is based on the use of molecular phylogenetic data. The current practice shows that some authors insist on considering panbiogeography as the only appropriate approach and vicariance as the only relevant process, whereas others accept Croizat's dictum "Earth and life evolve together" as a useful guide to understanding broad, general patterns, but recognize that dispersal also contributes substantially to biotic assembly. The framework of integrative pluralism allows to explain the complexities of the biogeographic processes involved in biotic assembly without the need of unification on a large scale. This historical analysis intersects with the existing historiography of the Modern Synthesis and may provide some insights on the dynamics of integrative pluralism, which may be especially relevant in the current development of the Extended Synthesis.

Understanding transmissibility patterns of Chagas disease through complex vector-host networks.

Chagas disease is one of the most important vector-borne zoonotic diseases in Latin America. Control strategies could be improved if transmissibility patterns of its aetiologic agent, Trypanosoma cruzi, were better understood. To understand transmissibility patterns of Chagas disease in Mexico, we inferred potential vectors and hosts of T. cruzi from geographic distributions of nine species of Triatominae and 396 wild mammal species, respectively. The most probable vectors and hosts of T. cruzi were represented in a Complex Inference Network, from which we formulated a predictive model and several associated hypotheses about the ecological epidemiology of Chagas disease. We compiled a list of confirmed mammal hosts to test our hypotheses. Our tests allowed us to predict the most important potential hosts of T. cruzi and to validate the model showing that the confirmed hosts were those predicted to be the most important hosts. We were also able to predict differences in the transmissibility of T. cruzi among triatomine species from spatial data. We hope our findings help drive efforts for future experimental studies.

[Diversity and biogeographic affinities of sharks, rays and chimaeras (Chondrichthyes: Elasmobranchii, Holocephali) of Mexico]. / Diversidad y afinidades biogeográficas de los tiburones, rayas y quimeras (Chrondrichthyes: Elasmobranchii, Holocephali) de México.

The diversity of chondrychthyans in Mexico is described. The fauna is composed by 214 species (111 sharks, 95 rays and 8 chimaeras) and represents 17.3 % of the total number of species recorded worldwide. The families with the highest diversity comprise: Rajidae (14.5 %), Carcharhinidae (12.1 %), Pentanchidae, Triakidae, and Urotrygonidae (5.1 %). In terms of geographical distribution, the diversity on the Mexican Pacific slope reaches up to 56.1 % of those species inhabiting Mexican marine and brackish waters (120 species, 62 genera, 37 families and 14 orders); the diversity in the Atlantic slope resulted similar to that on the Mexican Pacific with 55.1 % of the species (118 species, 59 genera, 35 families and 13 orders). The biogeographical affinities of the Mexican chondrychthyan fauna are complex with 19.7 % of the species being circumglobal, 9.9 % transatlantic, 1.9 % transpacific, and 9.4 % endemic to the exclusive economic zone. Additionally, 36.6 % of the species recorded so far are endemic to the Eastern Pacific coast where the species are similar to those found in the Cortez biogeographic province (27.7 %), followed by the Californian (20.7 %), Panamanian (19.3 %), Galapagos (5.6 %) and Peruvian-Chilean (8.9 %). Likewise, 33.3 % are endemic of the Atlantic coast, where species are similar to those found in the Caribbean province (31.9 %), followed by the Carolinean (24.4 %) and the Brazilian (6.6 %).

Cladistic biogeography of the Neotropical region: identifying the main events in the diversification of the terrestrial biota.

A cladistic biogeographical analysis was undertaken to identify the main events in the biotic diversification of the terrestrial Neotropical biota. For the 36 animal and plant taxa analysed, a component × area matrix was constructed, associating geographical data only with informative nodes, and it was analysed under implied weights using the software TNT. The general area cladogram obtained shows that the Neotropical region constitutes a monophyletic unit, with a first split separating the Antilles and a second one dividing the continental areas into a north-western and a south-eastern component. Within the north-western component the areas split following the sequence northern Amazonia, south-western Amazonia, north-western South America, and Mesoamerica. Within the south-eastern component the areas split following the sequence south-eastern Amazonia, Chaco, and Parana. The three main components are treated as subregions: Antillean, Amazonian (northern Amazonian, south-western Amazonian, Mesoamerican, and north-western South American dominions), and Chacoan (south-eastern Amazonian, Chacoan, and Parana dominions). Dispersal and vicariant events postulated to explain these pattens might have occurred during the Cretaceous, when the Caribbean plate collided with the Americas, a combination of eustatic sea-level changes and tectonic deformations of the continental platform exposed large parts of South America to episodes of marine transgressions, and the Andean uplift reconfigured the Amazonian area. Tertiary and Quaternary events are assumed to have later induced the diversification within these large biogeographical units.

Distributional patterns of Mawsoniidae (Sarcopterygii: Actinistia).

Mawsoniidae are a fossil family of actinistian fish popularly known as coelacanths, which are found in continental and marine paleoenvironments. The taxon is considered monophyletic, including five valid genera (Axelrodichthys, Chinlea, Diplurus, Mawsonia and Parnaibaia) and 11 genera with some taxonomical controversy (Alcoveria, Changxingia, Garnbergia, Heptanema, Indocoelacanthus, Libys, Lualabaea, Megalocoelacanthus, Moenkopia, Rhipis and Trachymetopon). The genera restricted to the Northern Hemisphere (Diplurus and Chinlea) possess the oldest records (Late Triassic), whereas those found in the Southern Hemisphere (Mawsonia, Axelrodichthys, and Parnaibaia) extend from Late Jurassic to Late Cretaceous, especially in Brazil and Africa. We identified distributional patterns of Mawsoniidae, applying the panbiogeographical method of track analysis, and obtained three generalized tracks (GTs): GT1 (Northeastern Newark) in strata of the Newark Group (Upper Triassic); GT2 (Midwestern Gondwana) in the Lualaba Formation (Upper Jurassic); and GT3 (Itapecuru-Alcântara-Santana) in the Itapecuru-Alcântara-Santana formations (Lower Cretaceous). The origin of Mawsoniidae can be dated to at least Late Triassic of Pangaea. The tectonic events related to the breakup of Pangaea and Gondwana and the evolution of the oceans are suggested as the vicariant events modeling the distribution of this taxon throughout the Mesozoic.

Biogeographical regionalisation of the Neotropical region.

A biogeographic regionalisation of the Neotropical region is proposed as a hierarchical classification of sub-regions, dominions, provinces and districts. This regionalisation is based on biogeographic analyses of terrestrial plant and animal taxa, and seeks to provide universality, objectivity and stability, such that it can be applied when describing distributional areas of particular taxa or comparing different biogeographic analyses. The Neotropical region is currently comprised of three sub-regions (Antillean, Brazilian and Chacoan), two transition zones (Mexican and South American), seven dominions (Mesoamerican, Pacific, Boreal Brazilian, Southwestern Amazonian, Southeastern Amazonian, Chacoan and Parana) and 53 provinces. For some of the latter, sub-provinces and districts are recognized. Complete synonymies and brief descriptions of the areas are provided, as well as the endemic taxa that diagnose the different provinces.

Niche conservatism and convergence in birds of three cenocrons in the Mexican Transition Zone.

Background: The niche conservatism hypothesis postulates that physiological and phylogenetic factors constrain species distributions, creating richness hotspots with older lineages in ancestral climatic conditions. Conversely, niche convergence occurs when species successfully disperse to novel environments, diversifying and resulting in areas with high phylogenetic clustering and endemism, low diversity, and lower clade age. The Mexican Transition Zone exhibits both patterns as its biotic assembly resulted from successive dispersal events of different biotic elements called cenocrons. We test the hypothesis that biogeographic transitionallity in the area is a product of niche conservatism in the Nearctic and Typical Neotropical cenocrons and niche convergence in the Mountain Mesoamerican cenocron. Methods: We split the avifauna into three species sets representing cenocrons (sets of taxa that share the same biogeographic history, constituting an identifiable subset within a biota by their common biotic origin and evolutionary history). Then, we correlated richness, endemism, phylogenetic diversity, number of nodes, and crowning age with environmental and topographic variables. These correlations were then compared with the predictions of niche conservatism versus niche convergence. We also detected areas of higher species density in environmental space and interpreted them as an environmental transition zone where birds' niches converge. Results: Our findings support the expected predictions on how niches evolved. Nearctic and Typical Neotropical species behaved as predicted by niche conservatism, whereas Mountain Mesoamerican species and the total of species correlations indicated niche convergence. We also detected distinct ecological and evolutionary characteristics of the cenocrons on a macroecological scale and the environmental conditions where the three cenocrons overlap in the Mesoamerican region.

Biotic assemblages of gelatinous zooplankton in the Gulf of Mexico and adjacent waters: An evolutionary biogeographic approach.

Gelatinous zooplankton constitutes a polyphyletic group with a convergent evolutionary history and poorly known biogeographical patterns. In the Gulf of Mexico, a region with complex geological, hydrological, and biotic histories, the study of this group has been limited to taxonomical and ecological aspects. In this study, we implemented a track analysis to identify distributional patterns of gelatinous zooplankton in the Gulf of Mexico and adjacent waters based on a dataset of 6067 occurrence records corresponding to Hydrozoa, Scyphozoa, Cubozoa, Ctenophora, Chaetognatha, Thaliacea, and Appendicularia. Information was compiled from the Global Biodiversity Facility Information (GBIF) and Ocean Biodiversity Information System (OBIS) databases and peer-reviewed literature. Individual tracks were constructed by joining the minimum distance between the occurrence localities of each taxon using a minimum spanning tree algorithm. We identified generalized tracks using parsimony analysis of endemicity with progressive character elimination (PAE-PCE). The areas where different generalized tracks overlapped were considered to represent panbiogeographical nodes. Seven generalized tracks (two with nested patterns) and six panbiogeographical nodes were recognized, mainly in neritic zones. The distributional patterns of gelatinous zooplankton allowed us to identify four biogeographic areas, supporting previously proposed biogeographic schemes. Gelatinous zooplankton in the Gulf of Mexico showed a convergent spatial distribution that can be explained by vicariant and dispersal events. The historical biogeography of the gelatinous biotas of the Gulf of Mexico has been little studied compared to ecological approaches, and the lack of integrative studies considering historical patterns is evident. This type of research is fundamental to understanding the evolutionary history of natural resources from a spatial perspective, identifying sites of biodiversity and endemism, and establishing a biogeographic baseline of the region for further studies.

List of the flea species of the State of Chihuahua, Mexico, with new records and a new species of Strepsylla Traub, 1950 (Siphonaptera: Ctenophthalmidae: Neopsyllinae: Phalacropsyllini).

A total of 45 flea species, assigned to 22 genera, are documented for the state of Chihuahua, Mexico. Fourteen records are new for the state, five species are recorded for the first time for the country and the state, and the number of species of Mexico is increased to 177 species. Chihuahua now has 25.4% of the total flea species registered for the country. A new species of the genus Strepsylla is described from specimens from the Flora and Fauna Protection Area Cerro del Mohinora, in the northwestern part of the Sierra Madre Occidental. Strepsylla mohinora sp. nov. is included in a published taxonomic key for the species of Strepsylla.

Phylogenetics of the tribe Phalacropsyllini (Siphonaptera: Ctenophthalmidae: Neopsyllinae) based on molecular and morphological evidence.

Six genera of the tribe Phalacropsyllini were analysed based on molecular (18S rDNA and 28S rDNA) and morphological characters, using maximum parsimony analyses and Bayesian inference. The results support the paraphyly of the tribe Phalacropsyllini, as shown in a previous study based only on molecular characters. Three main clades were identified: one includes the genera Neopsylla, Epitedia, Catallagia, and Delotelis; another includes Phalacropsylla and Strepsylla; and a third one includes only Meringis. In order to propose a phylogenetic classification we recognize three tribes, transferring Epitedia, Catallagia and Delotelis to Neopsyllini; keeping Phalacropsyllini only for Phalacropsylla and Strepsylla; and proposing the new tribe Meringini for Meringis.

THE NEOTROPICAL SPECIES OF SMALL CARPENTER BEES OF THE SUBGENUS CERATINA (ZADONTOMERUS) ASHMEAD, 1899 (HYMENOPTERA: APIDAE).

The Neotropical species of the subgenus Ceratina (Zadontomerus) Ashmead are revised. We recognize seven new species, giving a total of 10 species for the region: Ceratina (Zadontomerus) capitosa Smith, C. (Z.) ignara Cresson, C. (Z.) nautlana Cockerell, C. (Z.) kopili new species, C. (Z.) basaltica new species, C. sapphira new species, C. (Z.) indigovirens new species, C. (Z.) rehanae new species, C. (Z.) raquelitae new species, and C. (Z.) tepetlana new species. We propose the following synonymies: C. abdominalis Smith, C. tehuacana Strand, and C. parignara Cockerell under C. (Z.) ignara; and C. bakeri Smith and C. nigriventris Friese under C. (Z.) nautlana. Also, we describe the previously unknown male of C. capitosa, and provide a key to the species, diagnoses, descriptions and illustrations of the new species.

Toward a biogeographic regionalization of the Nearctic region: Area nomenclature and digital map.

We provide a preliminary nomenclatural proposal and a digital map of the Nearctic region, based on published regionalizations, especially Dice (1943), and applying the International Code of Area Nomenclature. The Nearctic region is comprised of three subregions (one of them with two dominions), one transition zone and 29 provinces. The Arctic subregion, in northern North America and Greenland, includes the Eskimoan, Hudsonian, Aleutian and Sitkan provinces. The Western subregion, in western North America, includes the Californian dominion, with the Californian and Oregonian provinces; and the Rocky Mountain dominion, including the Montanian, Saskatchewan, Palusian, Artemisian, Coloradan, Kansan, Mohavian, Navahonian, Sonoran, Chihuahuan, Comanche, and Baja California provinces. The Alleghany subregion, in eastern North America, includes the Illinoian, Canadian, Carolinian, Texan, Austroriparian, and Tamaulipan provinces. The Mexican Transition Zone, situated in the area of overlap with the Neotropical region, includes the Sierra Madre Occidental, Sierra Madre Oriental, Transmexican Volcanic Belt, Sierra Madre del Sur and Chiapas Highlands provinces.

Revision of the genus Cenophengus LeConte, 1881 (Coleoptera, Phengodidae), with the description of four new species, new geographic records and a new synonymy.

A taxonomic revision of the genus Cenophengus LeConte, 1881 (Coleoptera: Phengodidae) is provided, including new data on geographic ranges of the species. This is the first time this genus has been recorded for Belize and in Honduras. Four new species (C.gardunoi, C.saasil, C.tsiik and C.zuritai) are described and a new synonymy (C.guerrerensis, Zaragoza-Caballero, 1991 = C.major Wittmer, 1976) is established. The study includes a key to the 30 valid species, diagnoses, descriptions, photographs and distribution maps.

The Triatoma phyllosoma species group (Hemiptera: Reduviidae: Triatominae), vectors of Chagas disease: Diagnoses and a key to the species.

The Triatoma phyllosoma species group includes 17 species of kissing bugs, most of them implicated in the transmission of Chagas disease in the Americas. The species of this group are T. bassolsae Alejandre-Aguilar, Nogueda-Torres, Cortz-Jmenez, Jurberg, Galvo Carcavallo, 1999, T. brailovskyi Martnez, Carcavallo Pelaez, 1984, T. dimidiata (Latreille, 1811), T. gerstaeckeri (Stl, 1859), T. gomeznunezi Martnez, Carcavallo Juberg, 1994, T. hegneri Mazzotti, 1940, T. huehuetenanguensis Lima-Cordn, Monroy, Stevens, Rodas, Rodas, Dorn Justi, 2019, T. indictiva Neiva, 1912, T. longipennis Usinger, 1939, T. mazzottii Usinger, 1941, T. mexicana (Herrich-Schaeffer, 1848), T. mopan Dorn, Justi, Dale, Stevens, Galvo, Lima-Cordn Monroy, 2018, T. pallidipennis (Stl, 1872), T. phyllosoma (Burmeister, 1835), T. picturata Usinger, 1939, T. recurva (Stl, 1868), and T. sanguisuga (LeConte, 1855). The validity of some species of the group was uncertain, because of both cryptic species and hybrid occurrence. Species exhibiting these particularities were formerly classified in the T. dimidiata and T. phyllosoma complexes. Although we recognize the historical value of these species complexes, we do not recommend their further use. Instead, we recognize the T. phyllosoma species group here reviewed, considering the current knowledge of the systematics and reproductive behavior of the group. We implement the cohesion species concept, validating the species status of T. bassolsae, T. longipennis, T. mazzottii, T. pallidipennis, T. phyllosoma, and T. picturata. We also provide diagnoses, photographs and a taxonomic key including the recently described species.

An Endemic and Endangered New Species of the Lizard Liolaemus montanus Group from Southwestern Peru (Iguania: Liolaemidae), with a Key for the Species of the L. reichei Clade.

The southwestern Andes of Peru harbor a hidden taxonomic diversity of reptiles. We describe a new species of Liolaemus Wiegmann (Iguania: Liolaemidae) from xerophytic environments of the southwestern slopes of the Andes of Peru, 2,400-2,900 m asl. The new species, previously considered to be a population of L. insolitus Cei, exhibits unique diagnostic characters of morphology, scalation and color pattern, and molecular evidence that suggest that it belongs in the Liolaemus montanus species group and the L. reichei clade. Moreover, the species is endemic to the eastern slopes of La Caldera batholith in the Department of Arequipa, southern Peru. We also provide information on the conservation status of the species and suggest it be included in the IUCN red list of the threatened species as endangered (EN). A key for the species of the L. reichei clade is provided.