Biogeographic regions of North American mammals based on endemism

Since the 19th Century, two regions have been recognized for North American mammals, which overlap in Mexico. The Nearctic region corresponds to the northern areas and the Neotropical region corresponds to the southern ones. There are no recent regionalizations for these regions under the criterion of endemism. In the present study, we integrate two methods to regionalize North America, using species distribution models of mammals: endemicity analysis (EA) and parsimony analysis of endemicity (PAE). EA was used to obtain areas of endemism and PAE was used to hierarchize them. We found 76 consensus areas from 329 sets classified in 146 cladograms, and the strict consensus cladogram shows a basal polytomy with 14 areas and 16 clades. The final regionalization recognizes two regions (Nearctic and Neotropical) and a transition zone (Mexican Transition Zone), six subregions (Canadian, Alleghanian, Californian‐Rocky Mountain, Pacific Central America, Mexican Gulf‐Central America, and Central America), two dominions (Californian and Rocky Mountain), and 23 provinces. Our analysis show that North America is probably more complex than previously assumed. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 485–499.     

Regionalization of the avifauna of the Baja California Peninsula, Mexico: a parsimony analysis of endemicity and distributional modelling approach

Aim To analyse the distributional patterns of the Baja California Peninsula's resident avifauna, and to generate a regionalization based on a method that uses a parsimony analysis (parsimony analysis of endemicity, PAE) of point data and modelled potential distributions. Location The Baja California Peninsula, Mexico. Methods A data base was constructed containing records of 113 species of resident terrestrial birds present in the Baja California Peninsula. Records and localities were obtained from the literature and from specimens housed in scientific collections world‐wide. Raw data points and potential distribution maps obtained using the software Genetic Algorithms for Rule‐set Prediction (GARP), were analysed with PAE. Results The data base consisted of 4164 unique records (only one combination of species/locality) belonging to 113 terrestrial resident bird species, in a total of 809 localities. From the point distribution matrix, the analysis generated 500 equally parsimonious trees, from which a strict consensus cladogram with 967 steps was obtained. The cladogram shows a basal polytomy and some geographical correspondence of a few resolved groups obtained in the analysis. These results do not allow the recognition of areas defined by avifaunistic associations. From the potential distribution matrix, the analysis generated 501 equally parsimonious trees, and a strict consensus cladogram of 516 steps was obtained. The cladogram shows a higher resolution because of the number of resolved groups with better geographical correspondence and therefore regions are well‐defined. Main conclusions The correspondence of some groupings of species suggest their validity as areas with biogeographical (historical and/or ecological) meaning. This regionalization in the Baja California avifauna seems to be consistent with previous regionalizations for other groups. Hence, PAE is a useful tool for area categorization if reliable point records and prediction tools are available. Our results suggest that the geographical definition is much better using potential data generated by GARP, particularly when they are contrasted with the results from point data. Thus, this is an excellent alternative for developing biogeographical studies, as well as for improving the use of data from scientific collections and other sources of biodiversity information.     

Mexican Fruit Fly Populations in the Semi-Arid Highlands of the Sierra Madre Oriental in Northeastern Mexico

The Mexican fruit fly, Anastrepha ludens Loew (Diptera: Tephritidae), is one of the most important pests of citrus in Mexico. We report the results of an analysis of A. ludens populations that inhabit the semi-arid highlands of the Sierra Madre Oriental in northeastern Mexico. This study aimed to provide information on population fluctuation of A. ludens and how it relates to climate variables, as well as insights into habitat and native parasitoids. Population peaked in the period July–November when ripe fruits of the wild host, Casimiroa pubescens Ramírez, were available. No adults were captured the rest of the year, suggesting that high populations depend on the availability of wild host fruit. No significant relationships between population fluctuation and climatic variables were observed, except for minimum temperature. Fruit samples of citron (Citrus medica L.), pomegranate (Punica granatum L.), and C. pubescens were collected to determine degree of infestation. Infestation levels (pupae/g) ranged between 0.0006 for citron, 0.0047 for pomegranate, and 0.0240 for C. pubescens. A native parasitoid of Tephritidae, Doryctobracon crawfordii (Viereck) (Braconidae), was identified. Parasitism percentage was calculated at 12.5% on C. pubescens fruits. No parasitoids were observed on citron or pomegranate fruit samples. These results contribute to knowledge on behavior of A. ludens native to temperate environments where no commercial hosts are available. Further research on host expansion of this pest in light of scenarios of global climate change is suggested.     

Cladistic analysis of distributions and endemism (CADE): using raw distributions of birds to unravel the biogeography of the South American aridlands

Aim To use published distributional and taxonomic information on Neotropical birds in a cladistic analysis of distributions and endemism (CADE) to generate a testable hypothesis of area‐relationships for the arid areas of endemism, particularly those of Central South America (the ‘arid diagonal’), and to clarify the different methodologies commonly associated with parsimony analysis of endemicity (PAE). Location South America. Methods Cladistic analysis of distributions and endemism. Results We obtain a tree where the relationships of most areas are resolved, and we find support for an exclusive clade of Central South American areas, with the Caatinga as sister to both the Chaco and Cerrado. Main conclusions There is a substantial amount of historical signal in avian distributions, when large numbers of taxa and multiple taxonomic levels are considered. Ecological noise and historical information are more easily distinguished in CADE analyses than they would be in PAE analyses. Based on our results we predict that among aridland birds, the Cerrado and Chaco species will be more closely related to each other than to Caatinga species.     

中国斑腿蝗科特有种的分布及特有分布区划分

按照150km×150km对中国陆地部分进行栅格划分,收集整理了中国斑腿蝗科237个特有种在各栅格的分布信息,建立了物种(237)×栅格(168)数据矩阵,研究了中国斑腿蝗科特有种的分布规律并用PAE法进行特有分布区划分。结果表明:除天山山地分布的少数几个种外,中国斑腿蝗科特有种基本分布西南-东北走向。在该范围内南多北少。华南和西南是中国斑腿蝗科特有种最丰富的地区,在秦巴山区、藏东南和浙闽丘陵地带各有一个高密度的分布区。11个特有分布区中5个分布在东洋界部分,两个在中国东部古北和东洋界交界区,天山山地有一个分散的特有分布区,东北有2个特有分布区。特有分布区的分布和划分基本和中国动物地理区划及区系相一致,但台湾地区归入华中区;横断山和秦岭山地的栅格在支序图中多形成未决分类单元,特有分布区划分困难;华南和华中地区的特有分布区可以进一步划分成更小范围的特有分布区;从云南西北的腾冲、保山经过贵州南部到广西桂林的一个东西狭长的区域可以作为一个特有分布区     

Historical biogeography of the Yucatan Peninsula,Mexico: a perspective from ferns (Monilophyta) and lycopods (Lycophyta)

Based on known data sets and maximum entropy distribution data of fern and lycopod species registered in the Yucatán Peninsula, track and parsimony analyses were undertaken to evaluate the contribution of these groups to the establishment of biogeographical relationships of the peninsula with other areas. The resulting generalized tracks clearly agree with the geological origin of the peninsula and the previously recognized relationship with the Greater Antilles is not supported for ferns and lycopods. Instead, a Central American generalized track connects the Yucatán Peninsula with south‐eastern México and Central America. Floristically, the peninsula harbours 66 species of ferns and lycopods. Seven are registered for the first time in the Yucatán Peninsula and one is a new species for México. These species do not follow the latitudinal pattern expected if ecological factors, such as humidity and rainfall, were the most important in determining their distributions. Groups of areas recognized with parsimony analysis of endemicity could not be defined as provinces as a result of the lack of endemic species. Nevertheless, a regionalization scheme based on maximum entropy distribution data and supported by track analyses is proposed. Two separate districts are recognized within the Yucatán Peninsula: arid/dry Yucatán in the north and El Petén (humid) in the south. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 775–786.     

Historical biogeographical patterns of the species of Bursera (Burseraceae) and their taxonomic implications

Aim The plant genus Bursera, with 104 species of trees and shrubs, has been used as a model for biogeographical analyses because of its high species richness and large number of endemic species. The biogeographical patterns of Bursera and their implications for its phylogenetic classification are reviewed in order that some hypotheses on the historical biogeography of tropical Mexico can be proposed. Location Bursera is found in the south‐western USA, most of Mexico, mainly below 1700 m elevation in tropical forests, with some species in xeric shrublands, diversifying along the Pacific slope, Central America, and north‐western South America. A few species occur on the Galapagos and Revillagigedo archipelagos, some of which are endemics, whereas in the Antilles species are distributed extensively, with several endemics in the Bahamas, Cuba, Jamaica, and Hispaniola. Methods Data from specimens in herbaria and the literature were used to construct a matrix of 104 species in 160 areas. Distributional patterns of the species of Bursera were inferred applying track analysis, parsimony analysis of endemicity (PAE), and Brooks parsimony analysis (BPA). Results Track analysis revealed four individual tracks: (1) a circum‐Caribbean track, comprising species of the Bursera simaruba species group; (2) an Antillean track, including species that have been transferred to Commiphora based on their pollen traits; (3) a Mexican Pacific track, including species of the B. fragilis, B. microphylla, and B. fagaroides species groups, called ‘cuajiotes’; and (4) a Neotropical Pacific track, including the two species groups assigned to section Bullockia, in which the individual track of the Bursera copallifera species group is nested within the track of the B. glabrifolia species group. The four tracks overlap in a node in the Mexican Pacific slope, where they are highly diversified. PAE allowed us to identify 22 areas of endemism: 12 in Mexico (11 along the Mexican Pacific slope), six in the Antilles, two in Central America, one in South America, and one in the Galapagos. The general area cladogram obtained by BPA has two main clades: one includes the greater Antilles; and the other, 12 Mexican areas of endemism. Main conclusions Bursera fragilis, B. microphylla, and B. fagaroides species groups can be treated together as a new section within Bursera, sect. Quaxiotea, because they are segregated from the other groups of sect. Bursera based on morphological, anatomical, molecular and geographical evidence.     

Breeding biology of Eared Quetzals in the Sierra Madre Occidental, Mexico

ABSTRACT.   Eared Quetzals ( Euptilotis neoxenus ), a threatened species, are one of the least studied trogons in Mexico. We monitored 29 Eared Quetzal nests in the Chihuahuan portion of the Sierra Madre Occidental from 1998 to 2003. All nests were in tree cavities, and the mean tree and nest cavity heights ( N = 14) were 16.9 ± 7.8 m and 11.4 ± 4.1 m, respectively. The mean clutch size was 2.8 ± 0.9 eggs ( N = 28), the incubation period lasted 22 d ( N = 1), and nestling periods ranged from 29 to 31 d ( N = 5). Both adults incubated eggs and fed nestlings. Of 80 eggs, 70 hatched (87.5%) and 67 of 70 young fledged (95.7%). Twenty-five of 29 nests (86.2%) produced at least one fledgling. One nest was predated, and two failed when nest trees fell. Higher rates of nest predation have been reported for other species of trogons. However, fewer potential predators, such as snakes and mammals, are present in the Sierra Madre than in tropical zones where most trogon species occur. In addition, antipredator behaviors, including nestlings with calls resembling a snake and nests with an unpleasant odor, may contribute to the high nesting success. The main limiting factors for Eared Quetzals in the northern Chihuahua may be competition for cavities with other secondary cavity-nesters, and the failure of nests when snags fall.     

Biodiversity and biogeography of the avifauna of the Sierra Madre Occidental,Mexico

The Sierra Madre Occidental (SMOc) is located in the boundary between the Nearctic and Neotropical regions, area which has been considered as a complex transition zone. We analysed biogeographic patterns of its resident avifauna, including species richness, endemism, and biotic regionalization by analysing presence-absence matrices of 148 species of resident-terrestrial birds. We created the species richness maps by overlapping potential distribution maps obtained for each species via species distribution models (SDMs). To depict biogeographic patterns, we used strict consensus cladograms from parsimony analyses of endemicity (PAE) and phenograms from an unweighted pair-group method with arithmetic average clustering algorithm. The Pacific slope of the SMOc contains the highest species richness, decreasing towards the northeast, and reflected in endemic and endangered species richness patterns. The PAE resulted in one area of endemism represented by the whole SMOc, outlining a divided area in its Pacific slope. The cluster analyses divided the area into two. One group towards the Pacific slope, delimited by the mountain ridge and characterized by tropical vegetation types and Mexican-Mesoamerican affinities; the other group is located towards the east and northeast, characterized by arid and temperate types of vegetation and Nearctic affinities. These results evidence a transition from a tropical to a temperate composition of bird species. In this way the location for a boundary between the Nearctic and the transition zone, for birds in this part of Mexico, is restricted to these highest elevations.     

Parsimony analysis of endemicity (PAE) of Mexican terrestrial mammals at different area units: when size matters

Abstract Aim Parsimony analysis of endemicity (PAE) is a biogeographical method that uses a parsimony algorithm to obtain an area cladogram, based on taxa inhabiting the study areas. We compare its performance at different geographical units (½° and 1° quadrats, ecoregions and biogeographical provinces) to analyse distributional patterns of Mexican terrestrial mammals, in order to assess the importance of the size of area units. Location The area analysed corresponds to Mexico. Methods Parsimony analyses were based on 56,859 collection records, corresponding to 703 genera, species and subspecies. Four data matrices were constructed for: (1) 716 quadrats of ½° latitude × ½° longitude, (2) 230 quadrats of 1° latitude × 1° longitude, (3) forty‐five ecoregions and (4) fourteen biogeographical provinces. Results For the ½° quadrat matrix, we obtained six cladograms of 17,138 steps. For the 1° quadrat matrix, we obtained five cladograms (strict consensus with 9394 steps). For the matrix of ecoregions, we obtained twelve cladograms (strict consensus cladogram with 3009 steps). For the provinces, we obtained a single cladogram with 1603 steps. Main conclusions The best results were obtained with natural areas instead of quadrats. There seems to exist a trend to decrease the absolute number of steps and an increase in the absolute and relative number of synapomorphies as the size of the area units decreases, although this does not necessarily occur for the number of cladograms.     

Areas of endemism of Mexican mammals: reanalysis applying the optimality criterion

In order to test Mexican areas of endemism of mammals identified by previous parsimony analyses of endemicity (PAEs), we applied the optimality criterion to three data matrices (based on point records, potential distributional models and the fill option in software NDM). We modelled the ecological niches of 429 terrestrial mammal species using the genetic algorithm for rule-set prediction (GARP) and models were projected as potential distributional areas. We overlapped the point occurrence data and the individual maps of potential distributions to a grid of 1° latitude–longitude. Three matrices of 247 grid cells (areas) and 429 species were built: (1) a binary matrix with '0' for absence and '1' for presence of at least one record of the species inside the grid-cell; (2) a three-state matrix similar to (1) but assigning the state '2' to the assumed presence in the model of potential distribution; and (3) a three-state matrix similar to (2), but applying the fill option of software NDM instead of using a model. The optimality criterion was performed in NDM version 2.7 and results were examined with VNDM version 2.7. The first and second matrices showed 13 areas of endemism and the third identified 16 areas of endemism. NDM provided a better resolution than PAE, allowing us to identify several new areas of endemism, previously undetected. Ecological niche models, projected as potential distributional areas, and the optimality criterion are very useful to identify areas of endemism, although they should be used with caution because they may overpredict potential distributional areas. PAE seems to underestimate the areas of endemism identified.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 468–478.     

A new Miocene local fauna from the Sierra Madre Oriental at San Luis Potosí, Central-East Mexico,and its paleontologic significance

Mexico's Late Neogene mammal faunas are largely known from localities in the Trans-Mexican Volcanic Belt; those from other morphotectonic provinces are few and far apart. Thus, the discovery of Late Miocene vertebrates in western Sierra Madre Oriental at San Luis Potosí, the Paso del Águila local fauna, significantly adds to this meager record. The assemblage was collected from the floodplain facies of the San Nicolás Formation, a ∼1100-m thick, dominantly fluviolacustrine and calcilithitic, 15°–20° NE dipping sequence preserved in the Peotillos-Tolentino Graben, between 22°11’–22°19’ N and 100°30’–100°39° W. It includes remains of cf. Trachemys, a small to medium-sized emydid chelonian, a large camelid, a small cervid and a new species of the equini Pliohippus s.s., comparable in size, cranial morphology and odontographic characters to the Clarendonian-Early Hemphillian horses of the Pliohippus clade. Ar-Ar dates from ash-fall tuffs seemingly above and below the fossiliferous strata, bracket the age between 12.33 and 7.41 Ma (i.e., late Middle to Late Miocene), that is, within the Late Clarendonian-Early Hemphillian NALMA interval, making this fauna the first in Mexico from this age. The Paso del Águila local fauna is at least partly correlative with the Hemphillian local faunas from the TMVB and adjacent areas (e.g., Rancho El Ocote, Guanajuato and Tecolotlán, Jalisco), the Central Plateau (e.g., Arroyo Los Fragmentos, Zacatecas), and the Sierra Madre Occidental (e.g., Yepómera). Elsewhere, it is broadly correlative with the Late Clarendonian-Early Hemphillian faunas from the California Coast Ranges (e.g., North Tejon Hills, Ricardo and Dove Springs in the Mohave Desert), and the Gulf Coast Plain, Florida (McGehee Farm and Mixon). The Paso del Águila local fauna was part of a subtropical savannah and pine-oak forest (with a well-developed understory) biome that thrived on a climate regime much more humid than today.     

Application of parsimony analysis of endemicity in Amazonian biogeography: an example with primates

The distributions of 51 non-human primate species are used for Parsimony Analysis of Endemicity (PAE) to determine the relationships among 14 interfluvial regions in the Amazon basin, South America. Two most parsimonious cladograms were found. The strict consensus tree of these cladograms suggests an early separation between Lower Amazonia (eastern) and Upper Amazonia (western). The major clusters of interfluvial regions identified in the PAE cladogram are congruent with the areas of endemism delimited for birds. When interfluvial regions are converted into avian areas of endemism, the PAE cladogram is congruent with one of the two general areas cladograms suggested for Amazonia based on phylogenies of several clades of forest birds. Our analysis suggests that PAE can be used as a tool to objectively identify areas of endemism at an intra-continental scale as well as to make historical inferences. However, the value of a PAE cladogram in this latter application should be always evaluated by congruence with area cladograms built upon cladistic biogeography procedures.     

Spatial niche modelling of five endemic cacti from the Brazilian Caatinga: Past,present and future

Climate change, together with human activities, impacts on natural and human systems on all continents and poses a major threat to biodiversity, especially in environments with a high rate of endemism and where species are profoundly adapted to specific environmental conditions, as is the case of the seasonally dry tropical forests, noticeably the Caatinga, an exclusively Brazilian biome. The objective of this study was to build spatial niche models of five species of Cactaceae (Arrojadoa penicillata, Brasilicereus phaeacanthus, Pereskia aureiflora, Stephanocereus leucostele and Tacinga inamoena) endemic to the Caatinga and with different traits, to evaluate the impact of climate change on their geographical distribution. The species records and environmental variable values were overlaid on a grid of 6818 cells with 0.5° spatial resolution. Niche models were obtained for five types of general circulation models between ocean and atmosphere and 12 different ecological models. The ensemble ecological niche model was calculated at present and projected to past (last glacial maximum – LGM, 21 000; and mid‐Holocene – Hol, 6000 years ago) and future climate conditions (average of 2080), under the effect of climate change, in the greenhouse gas emission scenario RCP4.5. The distribution pattern of the studied species indicates an area with less environmental suitability in the LGM, followed by an expansion that began in the Hol and continued until the present period. In the future (2080), the models predicted a retraction of areas of environmental suitability, in which P. aureiflora and B. phaeacanthus, given their more restricted, marginal habitat and woody habit, present a great risk of extinction, whilst S. leucostele, A. penicillata and T. inamoena present a smaller reduction in suitable area, partly reflecting their spreading, less woody habit. Regional conservation actions for Cactaceae species and their habitat need to take these findings into account if we are to ensure the survival of these species.     

Historical biogeography of South American freshwater fishes

Aim To investigate biogeographical patterns of the obligate freshwater fish order Characiformes. Location South America. Methods Parsimony analysis of endemicity, likelihood analysis of congruent geographical distribution, and partition Bremer support were used. Results Areas of endemism are deduced from parsimony analysis of endemicity, and putative dispersal routes from a separate analysis of discordant patterns of distribution. Main conclusions Our results demonstrate the occurrence of 11 major areas of endemism and support a preferential eastern–western differentiation of the characiforms in the Amazonian region, contrasting with the southern–northern differentiation of terrestrial organisms. The areas of endemism identified seem to be deeply influenced by the distribution of the emerged land during the 100‐m marine highstand that occurred during the late Miocene and allow us to hypothesize the existence of eight aquatic freshwater refuges at that time. The raw distribution of non‐endemic species supports nine patterns of species distribution across the 11 areas of endemism, two of which support a southern–northern differentiation in the eastern part of the Amazon. This result shows that the main channel of the Amazon limited dispersal between tributaries from each bank of the river. The levels of endemism further demonstrate that the aquatic freshwater refuges promoted allopatric speciation and later allowed the colonization of the lowlands. By contrast, the biogeographical pattern found in the western part of the Amazon is identified as a result of the Miocene Andean foreland dynamic and the uplift of the palaeoarches that promoted allopatric divergence across several sedimentary basins by the establishment of disconnected floodplains. The assessment of conflicting species distributions also shows the presence of seven putative dispersal routes between the Amazon, Orinoco and Paraná rivers. Our findings suggest that, rather than there being a single predominant process, the establishment of the modern South American freshwater fish biotas is the result of an interaction between marine incursions, uplift of the palaeoarches, and historical connections allowing cross‐drainage dispersal.     

Track analysis of the Nearctic region: Identifying complex areas with mammals

The Nearctic region is located on the North American plate. However, its tectonic history is related to convergence with other plates, which has promoted a complex topography. This complexity should be reflected by the distributional patterns of the biota. We used track analysis with 574 species of mammals to identify generalized tracks and panbiogeographic nodes in the Nearctic region and to propose an updated point of view of complex areas and their boundaries in North America. Seven generalized tracks with nested patterns (California, Columbia Plateau, Mesoamerican, Mexican Plateau, Neotropic, Southern Rocky Mountains, and Western Coast of USA) were identified using a parsimony analysis of endemicity with progressive character elimination. Nine panbiogeographic nodes were identified at the intersections of the generalized tracks, all of which were located in the Sierra of Chiapas and Central America physical features. A total of 192 nodes were identified for the nested patterns, located in only eight physical features. Our analysis revealed evolutionary patterns in generalized tracks, and the panbiogeographic nodes predicted areas with high evolutionary–geologic complexity, shared by other taxonomic groups.     

Biogeographical analysis of the flea beetle genus Chaetocnema in the Afrotropical Region: distribution patterns and areas of endemism

Aim The areal distributions of Chaetocnema species in the Afrotropical Region have been analysed with the aims of determining the distribution patterns (chorotypes) and identifying the most important areas of endemism for this flea beetle genus in sub‐Saharan Africa. Location Data were collected in sub‐Saharan Africa, including Madagascar. Methods The Afrotropical Region was divided into 103 5° quadrats (operative geographical units, or OGUs). A presence–absence matrix of the Afrotropical Chaetocnema species in the OGUs was analysed by cluster analysis (Baroni Urbani & Buser index and the WPGMA clustering method) to generate distribution pattern data based on similarity of distribution. The most important areas of endemism were identified by parsimony analysis of endemicity. Results The general distribution of Chaetocnema in the Afrotropical Region was found to be associated with moist environments and montane grasslands. Most species exhibit restricted geographical ranges. Cluster analysis revealed 120 spatial distributions that can be grouped into 13 distinct distribution patterns (chorotypes). The most important areas of endemism for Chaetocnema in sub‐Saharan Africa according to the present parsimony analysis of endemicity are: (1) central and eastern Madagascar [endemicity rate (ER) = 61.1%], (2) Western Cape Province (ER = 36.4%), (3) southern Drakensberg (ER = 26.7%), (4) the Shaba Region (ER = 16.7%), and (5) the North‐Kivu Region (ER = 5.0%). Main conclusions There are 123 known species of Chaetocnema in the Afrotropical Region, more than in any other zoogeographical region. About 91% of the species are endemic and they generally exhibit a restricted and often very localized geographical range. The remaining 9% of the species are represented by seven species that also inhabit northern Africa and/or the Arabian peninsula (C. bilunulata Demaison, C. ganganensis Bechyné, C. ljuba Bechyné, C. pulla Chapuis, C. tarsalis Wollaston, and C. wollastoni Baly), three species that widely inhabit the Palaearctic Region (C. conducta (Motschulsky), C. schlaeflini (Stierlin), and C. tibialis (Illiger)), and two species that were introduced (C. confinis Crotch, and C. picipes Stephens).