The biogeography of Hoffmannseggia (Leguminosae, Caesalpinioideae, Caesalpinieae): a tale of many travels

Aim The flowering plant genus Hoffmannseggia consists of 21 species distributed amphitropically between the arid regions of the south‐western United States and adjacent Mexico, and west‐central South America. This pattern of geographical disjunction is shared by numerous other angiosperm genera and has been the subject of discussions for more than a century with various authors advocating a northern origin for particular taxa and others advocating a southern origin. This study uses a well‐supported phylogeny of a genus with numerous species in each area to address the issues of a northern or southern origin and the facility with which organisms move between the two continents. Location South‐western United States and northern Mexico, northern Chile and Argentina, southern Bolivia, and western Peru. Methods Using DNA sequence data from the nuclear and chloroplast genomes, we generated a phylogenetic hypothesis for all species of Hoffmannseggia rooted with Zuccagnia and Balsamocarpon. Geographical data were optimized on the resultant tree to assess the probable continent of origin for the genus, the pattern of disjunctions between North and South America, and species radiations within the genus. Main conclusions Hoffmannseggia arose in South America and initially split into a suffrutescent (somewhat woody) and an herbaceous clade. Within each of these major clades, there have been at least two exchanges between North and South America. There are no data to support an ancestral pan‐American range for Hoffmannseggia and we therefore ascribe the amphitropical disjunctions to long‐distance dispersal. The phylogeny clearly shows that all dispersals were from South to North America and they occurred at different times and thus the pattern is not the result of a single simultaneous set of dispersals.     

Early evolution of Nemopteridae illuminated with the first and oldest thread‐winged lacewing in Cretaceous amber

Nemopteridae are a charismatic family of lacewings distinguished by specialization of morphology in both larvae and adults as well as in their biology and behaviour. The evolutionary history of Nemopteridae is largely unknown due to the scarcity of fossil records. Here we report a new genus and species of Nemopteridae (Cretocroce xiai gen. et sp.n.) based on an exquisitely preserved adult specimen in mid‐Cretaceous Burmese amber. We place the new genus in the subfamily Crocinae, which is characterized by filamentous hindwings with a sensory function as an adaptation to their cavernicolous habitats. The new genus represents the first nemopterid in amber and the oldest crocine known to date. The new fossil crocine is nearly morphologically unchanged over 99 Ma compared with its modern counterparts. Moreover, it is estimated to be closely related to some extant crocines endemic to South America based on morphological phylogenetic analysis. The new finding provides direct evidence to demonstrate the antiquity of Crocinae morphological innovation and related biology, such as pollenophagous habits, and further corroborates Gondwanan origin of the subfamily. The published work has been registered on ZooBank: http://zoobank.org/urn:lsid:zoobank.org:pub:3D29DE48‐DBD8‐4614‐8387‐D9FCF90B0640 .     

Three new species of Crenitis Bedel, 1881 from South Africa,with a revised key to African species (Coleoptera: Hydrophilidae)

Three new species of Crenitis Bedel, 1881 are described from South Africa; C. castellus sp.n. and C. rupestris sp. n. from the KwaZulu-Natal Drakensberg, and C. quagga sp.n. from the Western Cape Province. All three new species were found in madicolous habitats, which appear to be characteristic for a number of species of the genus. A revised key to African species of Crenitis is included.

http://zoobank.org/urn:lsid:zoobank.org:pub:18BE876C-46CB-4AD5-9BB0-EA6F8D711F09     

Phylogeny and biogeography of the hollies (Ilex L., Aquifoliaceae)

The holly genus, Ilex L., in the monogeneric Aquifoliaceae, is the largest woody dioecious genus (>664 spp.), with a near‐cosmopolitan distribution in mesic environments. We constructed a phylogeny based on two nuclear genes, representing 177 species spread across the geographical range, and dated using macrofossil records. The five main clades had a common ancestor in the early Eocene, much earlier than previously suggested. Ilex originated in subtropical Asia and extant clades colonized South America by 30 Ma, North America by 23 Ma, Australia by 8 Ma, Europe by 6 Ma, and Africa by 4 Ma. South and North America were colonized multiple times. Ilex also reached Hawaii (10 Ma) and other oceanic islands. Macrofossil and pollen records show the genus has tracked mesic climates through time and space, and had a wider distribution before late Miocene global cooling. Our phylogeny provides a framework for studies in comparative ecology and evolution.     

Out of the Neotropics: newly discovered relictual species sheds light on the biogeographical history of spider ants (Leptomyrmex,Dolichoderinae, Formicidae)

Spider ants of the genus Leptomyrmex Mayr (Hymenoptera: Formicidae: Dolichoderinae) are conspicuous species of Australasian rainforests, with putative fossil relatives in the Neotropics and Europe. There is longstanding debate over the biogeographical history of the genus, with the Palaearctic and Neotropical regions proposed as alternate centres of origin. We propose a resolution of this debate with the recent discovery and analysis of an extant species from central Brazil, L. relictus sp.n. , which we describe from workers, males and brood. We sequence ten nuclear genes in the new species and in several Australian Leptomyrmex species, and append these data to a 54‐taxon, 10‐gene data matrix previously generated for the subfamily Dolichoderinae. We conduct phylogenetic and divergence dating analyses, and re‐evaluate the fossil record of the group. We recover Leptomyrmex relictus sp.n. as a member of the Leptomyrmex clade with high support. It is sister to the Australasian species, and the genus Leptomyrmex is, in turn, sister to a pair of Neotropical genera, Forelius and Dorymyrmex. We infer a Neotropical origin for the genus and estimate a mid‐Eocene (46 Ma, 95% CI 56 to 36 Ma) origin for the crown genus and an Oligocene origin for the Australasian clade (29 Ma, 95% CI 40 to 19 Ma). We confirm placement of the Dominican amber species ?L. neotropicus Baroni Urbani in the genus but reject a close relationship with the Palaearctic fossil taxa ?Leptomyrmula Emery and ?Usomyrma Dlussky, Radchenko & Dubovikoff, considering them incertae sedis in the subfamily (Dolichoderinae). In contrast to the mesophilic preferences of the Australasian species of Leptomyrmex, the new Brazilian species inhabits cerrado (dry savannah). Our results support a Neotropical origin for spider ants with dispersal to Australia. Rafting on west‐bound currents and/or a historical diversity imbalance between Australia and South America are proposed as alternate hypotheses to explain a pattern of biased E–W mid‐Tertiary dispersal for ants with austral distributions. This pattern is suggested by our results in conjunction with observations of other ant clades. Overall, our findings highlight the value of integrated taxonomy, critical interpretation of morphology, and a comparative phylogenetic framework when conducting palaeontological and biogeographical studies of insect species. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:6E9E6617‐6E53‐40B8‐82C7‐67F89A83C553 .     

Two new Amphipod families recorded in South America shed light on an old biogeographical enigma

The known diversity of freshwater amphipods in South America is substantially lower than on other continents. This has puzzled biologists for decades. Two hypotheses have been proposed in attempts to explain this pattern. According to the first one, the majority of amphipod lineages never dispersed across South America. The alternative hypothesis is that the recently diversified hyalellids have outcompeted and depleted the ancestral amphipod fauna. The recently discovered freshwater amphipod species Seborgia potiguar sp. nov. (Seborgidae) and Potiberaba porakuara gen. nov., sp. nov. (Mesogammaridae) from Brazil reveals the existence of two additional families of amphipods in South America. In the light of these discoveries we have analysed the amphipod faunistic structure of South America to test the above two biogeographic hypotheses. First, the number of amphipod families in South America is not as low as was thought. Falklandellididae are limited to the Falkland Islands and Chile. All other families (Ingolfiellidae, Bogidiellidae, Phreatogammaridae, Paraleptamphopidae, Pseudoingolfiellidae, Paracorpohiinae, Mesogammaridae and Seborgidae) but one (Dogielinotidae) share two properties: (1) they have a transoceanic distribution and (2) they are from subterranean waters. Since the dispersal ability of amphipods is limited, trans-oceanic disjunctions are best explained by plate tectonics, which implies their early origin, negating the first biogeographical hypothesis. These ancient families, for unknown reasons, survived only in a stable subterranean environment which can be regarded as a refuge. The only recent colonizer of the continent might be Dogielinotidae with the species-rich genus Hyalella. Although it cannot be determined whether hyallelids truly out-competed ancient amphipods, we suggest that the second hypothesis fits better to the data. Further findings of amphipods are expected in South America, especially from subterranean waters. This habitat is highly endangered in Brazil, and should be more rigorously protected. http://www.zoobank.org/urn:lsid:zoobank.org:pub:CBE21C1E-4748-4237-9EAD-FBD240A7D501     

Support for vicariant origins of the New Zealand Onychophora

Aim The distribution of Onychophora across the southern continents has long been considered the result of vicariance events. However, it has recently been hypothesized that New Zealand was completely inundated during the late Oligocene (25–22 Ma) and therefore that the entire biota is the result of long-distance dispersal. We tested this assumption using phylogenetic and molecular dating of DNA sequence data from Onychophora. Location New Zealand, Australia, South Africa, Chile (South America). Methods We obtained DNA sequence data from the nuclear genes 28S and 18S rRNA to reconstruct relationships among species of Peripatopsidae (Onychophora). We performed molecular dating under a Bayesian relaxed clock model with a range of prior distributions using the rifting of South America and South Africa as a calibration. Results Our phylogenetic trees revealed that the New Zealand genera Ooperipatellus and Peripatoides, together with selected Australian genera (Euperipatoides, Phallocephale and an undescribed genus from Tasmania), form a monophyletic group that is the sister group to genera from Chile (Metaperipatus) and South Africa (Peripatopsis and Opisthopatus). The relaxed clock dating analyses yielded mean divergence times from 71.3 to 78.9 Ma for the split of the New Zealand Peripatoides from their Australian sister taxa. The 0.95 Bayesian posterior intervals were very broad and ranged from 24.5 to 137.6 Ma depending on the prior assumptions. The mean divergence of the New Zealand species of Ooperipatellus from the Australian species Ooperipatellus insignis was estimated at between 39.9 and 46.2 Ma, with posterior intervals ranging from 9.5 to 91.6 Ma. Main conclusions The age of Peripatoides is consistent with long-term survival in New Zealand and implies that New Zealand was not completely submerged during the Oligocene. Ooperipatellus is less informative on the question of continuous land in the New Zealand region because we cannot exclude a post-Oligocene divergence. The great age of Peripatoides is consistent with a vicariant origin of this genus resulting from the rifting of New Zealand from the eastern margin of Gondwana and supports the assumptions of previous authors who considered the Onychophora to be a relict component of the New Zealand biota.     

New species of Nanomis Lugo-Ortiz and McCafferty, 1999 (Ephemeroptera: Baetidae) from Venezuela

As the first taxonomic and faunistic report of Nanomis Lugo-Ortiz and McCafferty, 1999 for Venezuela, we describe a new species, N. rasmusseni sp. n., based on material from Andean streams. Comparisons of the nymphal and imaginal stages of N. rasmusseni with those of N. galera Lugo-Ortiz and McCafferty, 1999 – the only species currently known for this genus – are done in detail. Further, additional morphological characters are discussed to delineate the genus from the other baetid genera. Likewise, ecological data and distribution of N. rasmusseni in the Andean highlands of Venezuela are provided, which improves knowledge of the richness and distribution of the genus in South America.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:C9DF3936-101F-4240-B067-AC5DF394C805     

Biogeography of ‘tropical Anagallis’ (Myrsinaceae) inferred from nuclear and plastid DNA sequence data

Aim ‘Tropical Anagallis’ corresponds to one of two evolutionary lineages within the genus Anagallis L. Generally, species within this lineage have a limited distribution in (sub‐)tropical regions in Africa or Madagascar. Two species, however, are endemic to South America, and exhibit a trans‐Atlantic disjunction with the rest of the species within the lineage. To investigate this disjunct distribution, as well as other dispersal events, the distribution of extant taxa was used to hypothesize the ancestral area(s) of distribution. Location Africa, Madagascar, Europe and South America. Methods Dispersal–vicariance analysis (DIVA) was used to optimize distribution areas onto parsimony and Bayesian phylogenies based on sequence data from four chloroplast loci and the nuclear internal transcribed spacers (ITS). Results Parsimony analysis gave one most parsimonious tree while Bayesian analysis resulted in a collapsed node due to alternative placements of Anagallis nummularifolia Baker, endemic to Madagascar. Optimization of the present distribution using DIVA, and the most parsimonious tree and six alternative topologies of the Bayesian analysis, show an origin of the lineage in Europe as most likely, although one topology indicates a broader ancestral distribution area. Dispersal to Africa appears to have been a single event, while two parallel dispersal events seem to have resulted in the American as well as Madagascan distributions. Main conclusions The lineage ‘tropical Anagallis’ evolved in Europe and may have been present in the Eocene boreotropical forests, although scarcity of fossils makes assessment of age difficult. Dispersal to South America is proposed to have been via the North Atlantic land bridge, or, more likely, through transport by the North Equatorial Current. Dispersal from Europe to Africa represents a single event, while dispersal to Madagascar from mainland Africa has occurred twice.     

Molecular phylogenetics of mouse opossums: new findings on the phylogeny of Thylamys (Didelphimorphia,Didelphidae)

The mouse opossums of the genus Thylamys constitute a group of species mainly adapted to open xeric‐like habitats and restricted to the southern portion of South America. We used molecular data (mitochondrial and nuclear sequences) to evaluate the phylogenetic and biogeographical relationships of all currently known living species of the genus, recognizing a new taxon from the middle and high elevations of the Peruvian Andes and evaluating the phylogenetic structuring within T. pallidior and T. elegans, as well as the validity of T. sponsorius, T. cinderella and T. tatei, and the haplogroups recognized within T. pusillus. Our results confirm the monophyly of the genus and that the Caatinga and the Cerrado inhabitants Thylamys karimii and T. velutinus are the most basal species in the radiation of Thylamys. We also calibrated a molecular clock which hypothesized a time of origin of the genus of about 24 My, with most species differentiating in middle and late Miocene and Plio‐Pleistocene times of South America.     

Reclustering the cluster flies (Diptera: Oestroidea,Polleniidae)

A phylogenetic analysis of selected oestroid taxa based on 66 morphological traits and sequences from three nuclear protein‐coding genes (CAD, MAC, MCS) resolved the composition and phylogenetic position of the former subfamily Polleniinae of the Calliphoridae – here resurrected at family rank as Polleniidae Brauer & Bergenstamm, 1889 stat. rev. Six species are transferred from the family Rhinophoridae to the Polleniidae: the Palaearctic genus Alvamaja Rognes, along with its single species Alvamaja chlorometallica Rognes, and five Afrotropical species comprising the carinata‐group formerly in the genus Phyto Robineau‐Desvoidy but here assigned to genus Morinia Robineau‐Desvoidy, i.e. M. carinata (Pape, 1987) comb.n. , M. lactineala (Pape, 1997) comb.n. , M. longirostris (Crosskey, 1977) comb.n. , M. royi (Pape, 1997) comb.n. and M. stuckenbergi (Crosskey, 1977) comb.n. The Polleniidae are monophyletic and, in agreement with most recent phylogenetic reconstructions, sister to the Tachinidae. The female of A. chlorometallica and a new species of Morinia of the carinata‐group (Morinia tsitsikamma sp.n. from South Africa) are described. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:25B0C220‐DEE4‐4B0C‐88EA‐35FDE298EBC5 .     

A tale of the nearly tail‐less: the effects of Plio‐Pleistocene climate change on the diversification of the African avian genus Sylvietta

Sylvietta is a broadly distributed group of African species inhabiting a wide range of habitats and presents an interesting opportunity to investigate the historic mechanisms that have impacted the biogeography of African avian species. We collected sequence data from 50 individuals and used model‐based phylogenetic methods, molecular divergence estimates and ancestral area estimates to construct a time‐calibrated phylogeny and estimation of biogeographic history. We estimate a southern African origin for Sylvietta, with an initial divergence splitting the genus into two clades. The first consists of arid‐adapted species, with a southern African origin and subsequent diversification north into Ethiopia–Somalia. The second clade is estimated as having a Congolian forest origin with an eastward pattern of colonization and diversification as a result of Plio‐Pleistocene forest dynamics. Additionally, two members of the genus Sylvietta display interesting patterns of intraspecific diversification. Sylvietta rufescens is an arid‐adapted species inhabiting southern Africa, and we recover two subclades with a divergence dating to the Pleistocene, a unique pattern for avian species which may be explained via isolation in arid habitat fragments in the early Pleistocene. Second, Sylvietta virens, a species endemic to Afro‐tropical forests, is recovered with geographically structured genetic diversification across its broad range, an interesting result given that recent investigations of several avian forest species have found similar and substantial geographically structured genetic diversity relating to Plio‐Pleistocene forest fragmentation. Overall, Plio‐Pleistocene habitat cycling played a significant role in driving diversification in Sylvietta, and this investigation highlights the substantial impact of climate‐driven habitat dynamics on the history of sub‐Saharan species.     

A new species and first record of Androdeloscia (Oniscidea: Philosciidae) from Colombia

The genus Androdeloscia includes 24 Neotropical species distributed in Central and South America. This study describes the new species Androdeloscia colombiana sp. nov. from El Chorro, Soplaviento, and from the Botanical Garden Guillermo Piñeres, Turbaco, Bolívar, Colombia. This is the first record of the genus in Colombia.

http://zoobank.org/urn:lsid:zoobank.org:pub:AEA56E26-1F6D-4DDA-BA02-D73DE9FAD198     

Glacidorbidae (Gastropoda: Heterobranchia) in South America: revision and description of a new genus and three new species from Patagonia

The Glacidorbidae, a family restricted to the Gondwanan realm (Tasmania, southeastern and southwestern Australia, and southern Argentina and Chile), previously included five genera with 20 identified species; 19 of them are Australian, with one genus and species, Gondwanorbis magallanicus (Meier-Brook & Smith, 1976 Meier-Brook, K. & Smith, B.J. (1976) Glacidorbis Iredale, 1943, a genus of freshwater prosobranchs with a Tasmanian-Southeast Australian-South Andean distribution. Archive für Molluskenkunde 106, 191–198. [Google Scholar]), from South America. Here we describe two new species of Gondwanorbis: Gondwanorbis fueguensis n. sp. from the freshwater gastropods province of Southern Patagonia (Argentina) and Gondwanorbis tricarinatus n. sp. from Chile, and a new genus and species from the freshwater gastropods province of northern Patagonia (Argentina), Patagonorbis nahuelhuapensis n. sp and n. gen.

http://www./zoobank.org/urn:lsid:zoobank.org:pub:62EA0972-3AEF-4188-8E6D-F10895CE2BEF     

Biogeography of the grasses (Poaceae): a phylogenetic approach to reveal evolutionary history in geographical space and geological time

The grasses (Poaceae) are the fifth most diverse family of angiosperms, including 800 genera and more than 10 000 species. Few phylogenetic studies have tried to investigate palaeo‐biogeographical and palaeo‐ecological scenarios that may have led to present‐day distribution and diversity of grasses at the family level. We produced a dated phylogenetic tree based on combined plastid DNA sequences and a comprehensive sample of Poaceae. Furthermore, we produced an additional tree using a supermatrix of morphological and molecular data that included all 800 grass genera so that ancestral biogeography and ecological habitats could be inferred. We used a likelihood‐based method, which allows the estimation of ancestral polymorphism in both biogeographical and ecological analyses for large data sets. The origin of Poaceae was retrieved as African and shade adapted. The crown node of the BEP + PACCMAD clade was dated at 57 Mya, in the early Eocene. Grasses dispersed to all continents by approximately 60 million years after their Gondwanan origin in the late Cretaceous. PACCMAD taxa adapted to open habitats as early as the late Eocene, a date consistent with recent phytolith fossil data for North America. C₄ photosynthesis first originated in Africa, at least for Chloridoideae in the Eocene at c. 30 Mya. The BEP clade members adapted to open habitats later than PACCMAD members; this was inferred to occur in Eurasia in the Oligocene. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 543–557.     

Soil tardigrade biodiversity with the description of a new eutardigrade genus and its phylogenetic position

Sixteen tardigrade species have been identified from a total of 943 specimens isolated from 69 positive soil samples collected in southern Spain (Andalusia, Huelva). Three genera (Hexapodibius, Xerobiotus and the new genus Sarascon) and eight species are new records for the Iberian Peninsula, increasing Iberian tardigrade biodiversity by 6%, and 10 species are new records for soil habitats, increasing soil tardigrade biodiversity by 9% (from 96 to 108 species). A newly discovered species and genus are described, followed by an analysis and discussion of the species' phylogenetic position and synapomorphies, based on morphological and total evidence (morphology and molecular – 18S and 28S rRNA – data combined) phylogenetic analyses. The new genus is distinguished from other Itaquasconinae genera, to which it is phylogenetically related, by having a Parascon buccopharyngeal apparatus and Ramajendas claws. In contrast to general eutardigrade evolution, which is primarily determined by claw morphology, itaquasconid evolution is more related to buccopharyngeal apparatus morphology, which shows homoplastic evolution at superfamily, family and subfamily levels, and also defines functional trophic groups in soil-related environments.

http://zoobank.org/urn:lsid:zoobank.org:pub:068B9D87-27E2-4B26-8920-FF03CEF9869A     

World travellers: phylogeny and biogeography of the butterfly genus Leptotes (Lepidoptera: Lycaenidae)

Leptotina butterflies (Lycaenidae, Polyommatiinae) are found mostly in tropical and subtropical areas around the globe, marginally penetrating into temperate regions. Here, we investigated phylogenetic and biogeographical relationships of most representatives of the subtribe, using both likelihood and Bayesian approaches. We also estimated the timing of their diversification. And lastly, we studied phylogeographic patterns of the most widespread species, Leptotes pirithous. DNA sequences from two mitochondrial (COI, COII) and two nuclear genes (wingless, Ef1α) were analysed for 13 species of the genus Leptotes Scudder and one species of the genus Cyclyrius Butler. Both genera together form a monophyletic clade, and Cyclyrius is rooted deep inside Leptotes. Therefore, we designate Cyclyrius to be a junior synonym of Leptotes. According to our study, the genus Leptotes originated between the late Eocene and early Oligocene (35–31 Ma). During the Miocene it dispersed to the rest of the southern hemisphere, with further speciation events within the Indo‐Australian region, and separate radiations in the Americas and the Afrotropics. Leptotes webbianus from the Canary Islands turned out to be sister to the American clade from which it split c. 12 Ma. Leptotes pirithous originated in Madagascar c. 4 Ma and invaded the whole of Africa and southern Europe, including numerous surrounding islands. Populations of L. pirithous from Mauritius and Madagascar turned out to represent a distinct species (Leptotes durrelli sp.n. ) and the same applies to the Australasian populations of Leptotes plinius (Leptotes lybas stat. rev. ). This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:20308930‐988B‐4327‐A35F‐CC983D46263B .     

Succinapion telnovi n. gen. et n. sp. of the tribe Kalcapiini (Coleoptera: Brentidae: Apioninae) in Baltic amber

A new genus and species, Succinapion telnovi n. gen. et n. sp. (Coleoptera: Curculionoidea: Brentidae: Apioninae: Kalcapiini) is described and illustrated from Upper Eocene Baltic amber. The new genus is similar to the genus Melanapion Wagner, 1930 but differs from it in having femora ventrally with spine at distal 1/3, simple claws, a longer rostrum, elytra weakly widened towards apex, longer antennae and slightly narrower elytral striae.http://www.zoobank.org/urn:lsid:zoobank.org:pub:2A95C49D-5589-4ACA-8A87-0DDF635BA25E