A new species of Didymodon Hedw. (Pottiaceae, Bryophyta) from Peru

Didymodon incurvus J. A. Jiménez & M. J. Cano, is described and illustrated as a new species based on specimens collected from the Puno department in southern Peru. The new taxon is distinguished from others species of the genus by its oblong-lanceolate and appressed leaves when dry, incurved margins above midleaf, and the presence of a bulging ventral costal pad of cells. A key to species of Didymodon in Peru is provided. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 221–226.     

Integrative taxonomy reveals cryptic Amazonian species of Pristimantis (Anura: Strabomantidae)

Hypotheses on the taxonomic status of two Bolivian Pristimantis with taxonomic problems are assessed by an integrative taxonomic approach that integrates three independent lines of evidence: external morphology, prezygotic reproductive barriers (advertisement calls) and reciprocal monophyly (phylogenetic analyses of partial 16S mtDNA sequences). Central Andean Bolivian populations previously assigned to either P. peruvianus or P. dundeei, and lowland Amazonian populations from southern Peru and northern Bolivia previously considered P. peruvianus do not correspond to these species. Indeed, multivariate analyses of qualitative and quantitative morphological and bioacoustic characters, and phylogenetic analyses support the hypothesis that they represent different, previously unknown, cryptic lineages. They are herein described as new species. The former is a sibling species of P. fenestratus that inhabits the Amazonian and semideciduous forests of the Andean foothills in central Bolivia. The latter is sibling to the Andean species P. danae and is parapatric to it in the Amazonian lowland forests and adjacent foothills of northern Bolivia, southern Peru and adjacent Brazil. Most species of Neotropical frogs, and especially Pristimantis, have been described by using external qualitative morphological characters only. An extended integrative taxonomic approach, as exemplified herein, may lead to the discovery of many other cryptic and sibling lineages that would increase the species numbers of tropical areas. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 97–122.     

Assessing the taxonomic status of tropical frogs through bioacoustics: geographical variation in the advertisement calls in the Eleutherodactylus discoidalis species group (Anura)

The taxonomic status of a very poorly known group of Andean frogs (the Eleutherodactylus discoidalis group) is assessed through acoustic and statistical analyses of differences in temporal parameters of advertisement calls, such as the number of pulses and the call duration, and also in a spectral parameter, dominant frequency. As these species are usually misidentified or ignored because of their taxonomic complexity in both ecological and biodiversity studies, we provide a bioacoustical diagnosis for each species in order to facilitate identification in the field. Differences in acoustic parameters support the specific status of Eleutherodactylus cruralis , E. discoidalis , Eleutherodactylus ibischi , and Eleutherodactylus madidi . The name E. cruralis is probably applied to three different species: the nominal form from Amazonian forests of the Andean slopes and adjacent lowlands, and two cryptic species restricted to inter-Andean dry valleys and cloud forests from central Bolivia. Moreover, the distribution of energy through the call and the aggregation of pulses seem to be useful meristic characters for detecting interspecific differences. Populations from each macrohabitat can be recognized by distinctive advertisement calls, usually corresponding to a recognized species. For the whole group, pulse rate is significantly correlated to latitude, which could indicate a speciation process along the Andes in relation to habitat changes and isolation. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 353–365.     

Analysis of Andes frogs (Phrynopus, Leptodactylidae, Anura) phylogeny based on 12S and 16S mitochondrial rDNA sequences

South American leptodactylid frogs of the genus Phrynopus occur in cloud-forest, páramo, subpáramo and puna habitats (1000–4400 m elevation) from Colombia to Bolivia. Currently, there are 34 described species; however, many additional species new to science have been reported from Colombia, Peru, and Bolivia. The phylogeny of the species-diverse Phrynopus is unknown and the position of the genus within Leptodactylidae is poorly understood. We present the results of a phylogenetic study based on 12S and 16S mitochondrial rDNA. Fifteen species of Phrynopus from Bolivia to Ecuador are included, along with several other genera of Leptodactylidae and representatives of other frog families. Our results indicate that Phrynopus is phylogenetically nested within Eleutherodactylus , whereas Phyllonastes is phylogenetically nested within Phrynopus . Based on the recovered phylogeny, we transfer Phrynopus simonsii to Eleutherodactylus , and show that Phrynopus carpish needs to be removed from Phrynopus . Eleven of the 13 Phrynopus species occurring in Central Peru share the absence of an external and internal ear.     

Morphological phylogenetics of Puya subgenus Puya (Bromeliaceae)

Puya , a large genus mostly from South America, has been taxonomically divided into two subgenera: Puyopsis and Puya. The latter includes only eight species distributed mainly in Chile, extending to Argentina, Bolivia, and Peru. The species of subgenus Puya are recognized by the presence of a sterile apex of the inflorescence branches, whereas those of subgenus Puyopsis have fertile flowers all along the branches. The objectives of this article were to determine whether this diagnostic character was synapomorphic for subgenus Puya , and to explore the relationships between its species. Parsimony analyses were performed for 43 taxa and 93 morphological characters, 87 of which were discrete and six continuous. In the analysis that included all characters, a single most parsimonious tree was found that supported subgenus Puya by two synapomorphic character states, including the diagnostic character of a sterile inflorescence branch apex and a blooming pattern in which flowers open gradually from base to apex. The trees were better supported when the continuous characters were included. Further studies are suggested to resolve the infrageneric classification of Puya and the relationships of the species belonging to subgenus Puya .  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 93–110.     

A new species of Halocnemum M.Bieb. (Amaranthaceae) from southern Turkey

Halocnemum yurdakulolii Yaprak is described as the second species of the previously monotypic genus Halocnemum. The species is endemic to the Göksu Delta in southern Turkey. The main morphological characteristics that separate H. yurdakulolii from H. strobilaceum (Pall.) M.Bieb. are growth form and spike morphology. Apart from these morphological differences, the species show a clear genetic differentiation. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 716–721.     

Late Oligocene mesotheriids (Mammalia, Notoungulata) from Salla and Lacayani (Bolivia): implications for basal mesotheriid phylogeny and distribution

The oldest known mesotheriids are referred to the genus Trachytherus , a taxon traditionally included in its own subfamily 'Trachytheriinae'. The ontogeny, morphological variation and systematics of the genus Trachytherus are still poorly understood and the monophyly of this taxon is questionable. For the first time, a large collection of basal mesotheriids (belonging to the genus Trachytherus ) yielded by the Bolivian Salla beds (Deseadan, late Oligocene) and previously referred to T. spegazzinianus was included in a large-scale analysis. This has led to the recognition of a unique new species in the locality of Salla-Luribay, distinct from T. spegazzinianus : Trachytherus alloxus sp. nov . Skulls of this species are described and cheekteeth variation range throughout ontogeny is discussed. These new data are incorporated in a parsimony analysis of basal mesotheriids. The most probable hypothesis of a clade T. spegazzinianus -mesotheriines is discussed. The distinction of the new Salla species from the Patagonian species T. spegazzinianus confirms the faunal differences observed between these Bolivian and Patagonian Deseadan localities. Moreover, the assignment to Trachytherus spegazzinianus of mesotheriid remains from the Bolivian Lacayani deposits complicates the problem of these faunal differences. Further systematic investigations on the Lacayani fauna and on the age of the Patagonian localities are needed.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 153–200.     

Introducing a new species of the fern genus Doryopteris (Pteridaceae,Polypodiopsida) from the Neotropics

Doryopteris adornata is described as a new species from west‐central South America (Brazil, Bolivia, Paraguay, and Argentina). The dense indument of scales on the abaxial surface of the lamina is the most prominent diagnostic feature of this new species. Illustrations of the diagnostic characters are provided, as well as a table comparing the morphological characters of D. adornata with those of the two most similar species. The ecology and distribution are described. The conservation status for the new species using World Conservation Union (IUCN) guidelines/criteria is recommended. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 73–77.     

Systematics and biogeography of the widespread Neotropical gekkonid genus Thecadactylus (Squamata), with the description of a new cryptic species

The systematics of the widespread neotropical gekkonid genus Thecadactylus is investigated using a combination of morphological, morphometric and mtDNA (cytochrome b ) sequence data, forming a total evidence dataset. The analysis tackles the common yet complex problem of a widespread taxon consisting of one or more cryptic species that are difficult to diagnose using morphology alone. The data are analysed using both maximum parsimony and Bayesian inference. Virtually all analyses resolve a well-supported south-western Amazonian clade distinct from the remainder of the recognized T. rapicauda . The south-western Amazonian clade is not only robustly supported, but also exclusive, geographically coherent and sufficiently distinct to warrant specific recognition. The new species is diagnosable on the basis of molecular sequences that are 23.0–26.9% divergent from those of T. rapicauda , and morphological evidence. Bayesian inference analysis robustly resolves meaningful and repeatable patterns of relationship. The biogeography of Thecadactylus is interpreted in the context of its two constituent species, and difficulties of resolving systematic and biogeographical patterns in widespread, cosmopolitan taxa are discussed. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 339–370.     

A revision of Tecoma Juss. (Bignoniaceae) in Bolivia

A revision of the genus Tecoma in Bolivia based on the author's herbarium research and extensive field observations is presented. Species delimitation is re-evaluated, resulting in the recognition of only four species, one of which, T. beckii J. R. I. Wood, is described as new. Tecoma sambucifolia is treated as var. sambucifolia (Kunth) J. R. I. Wood of the widespread and variable T. stans (L.) Juss., as it can only be distinguished on the basis of minor morphological characters and is not as geographically restricted as has been supposed. Tecoma fulva (Cav.) D. Don is treated as a variable species extending from central Peru to Chile and northern Argentina with six subspecies: ssp. fulva , ssp. guarume (A. DC.) J. R. I. Wood, ssp. tanaeciiflora (Kränzlin) J. R. I. Wood, ssp. arequipensis (Sprague) J. R. I. Wood, ssp. garrocha (Hieron.) J. R. I. Wood, and ssp. altoandina J. R. I. Wood, the latter described as new. The differences between T. tenuiflora (A. DC.) Fabris and the Peruvian T. rosifolia Kunth are discussed. Hybrids are formally recognized in the genus for the first time, and the implications for the accepted corolla morphology and pollination syndromes are discussed. Maps illustrate the geographical distributions of the species. All taxa discussed in the paper are illustrated.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 143–172.     

Markedly discordant mitochondrial DNA and allozyme phylogenies of tube-nosed fruit bats, Nyctimene, at the Australian–Oriental biogeographical interface

Australo-Papuan tube-nosed fruit bats of the genus Nyctimene reach their most westerly distribution on the island archipelagos of eastern Indonesia. A recent morphological examination indicates three species occur on Moluccan islands. Both allozyme electrophoresis and mitochondrial DNA (mtDNA) sequencing reveal there are only two species present, Nyctimene cephalotes and Nyctimene albiventer , but there is considerable disparity between the results obtained from the two genetic data sets. Allozyme data indicate N. albiventer occurs on Wokam, which sits on the Australian continental shelf and was joined to Australo-Papua during the last glacial maximum, and N. cephalotes on the other Moluccan islands, all of which are off the continental shelf. Divergence of these two species is dated at approximately 1.5 Mya. By contrast, the mtDNA gene tree shows two deep clades, one containing all specimens from Wokam and Yamdena, and the second all the specimens from the other islands. This especially marked incongruence between the two genetic data sets is ascribed to either a single introgression event of N. albiventer mtDNA into N. cephalotes on Yamdena at around 100 000 ya or lineage sorting of very old mtDNA lineages that coalesce a rather long time before the speciation event. These results highlight that caution should be exercised when relying on mtDNA as barcodes in species taxonomy. The continental-associated N. albiventer on Wokam has higher levels of allozyme heterozygosity and mtDNA nucleotide diversity than the N. cephalotes populations occurring on the more remote islands, indicating that this colonizing species has experienced bottlenecks and/or a low effective population size since speciation.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 589–602.     

Two new South American species of Struthanthus (Loranthaceae) posing a challenge to circumscription of neotropical genera

Two new, closely related species of Struthanthus (Loranthaceae) are described, S. meridionalis from southern Bolivia and S. prancei from northern Brazil, each showing significant deviations from the generic norm in their inflorescence morphology. In both cases, a very high proportion of inflorescences bear bracteolate and/or ebracteolate monads as lateral units rather than the triads which characterize the rest of the genus. The place of inflorescence morphology in the development of generic concepts in small-flowered neotropical Loranthaceae is briefly reviewed, leaving the two new species in Struthanthus for the time being.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 469–474.     

Skull anatomy of the Oligocene toothed mysticete Aetioceus weltoni (Mammalia; Cetacea): implications for mysticete evolution and functional anatomy

Toothed mysticetes of the family Aetiocetidae from Oligocene rocks of the North Pacific play a key role in interpretations of cetacean evolution because they are transitional in grade between dorudontine archaeocetes and edentulous mysticetes. The holotype skull of Aetiocetus weltoni from the late Oligocene (28–24 Ma) of Oregon, USA, has been further prepared, revealing additional morphological features of the basicranium, rostrum and dentary that have important implications for mysticete evolution and functional anatomy. The palate of Aetiocetus weltoni preserves diminutive lateral palatal foramina and associated delicate sulci which appear to be homologous with the prominent palatal foramina and sulci that occur along the lateral portion of the palate in extant mysticetes. In modern baleen whales these foramina allow passage of branches of the superior alveolar artery, which supplies blood to the epithelia of the developing baleen racks. As homologous structures, the lateral palatal foramina of A. weltoni suggest that baleen was present in this Oligocene toothed mysticete. Cladistic analysis of 46 cranial and dental characters supports monophyly of the Aetiocetidae, with toothed mysticetes Janjucetus and Mammalodon positioned as successive sister taxa. Morawanacetus is the earliest diverging aetiocetid with Chonecetus as sister taxon to Aetiocetus species. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 308–352.     

The anatomy of Odobenocetops (Delphinoidea, Mammalia), the walrus-like dolphin from the Pliocene of Peru and its palaeobiological implications

The Pliocene Beds of the Pisco Formation on the southern coast of Peru have yielded three new skulls of the walrus-like odontocete, Odobenocetops (Delphinoidea, Cetacea). Two of the skulls are from a slightly younger horizon than the holotype of O. peruvianus and belong to a different species, O. leptodon . The holotype of O. leptodon bears a 135 cm long needle-like right tusk and a small 25 cm long left tusk, the apex of which was erupted. The third skull, referred to a female of O. peruvianus , bears two small tusks similar in size to the small left tusk of the holotype. The new specimens include periotic, tympanic and ear ossicles, which confirm referral to the Order Cetacea and the morphology of the humerus reinforces affinities of odobenocetopsids to monodontids. Because the anterodorsal edge of its orbit is slightly concave O. leptodon had reduced anterodorsal binocular vision, a condition compensated for by the probable presence of a small melon (and inferred echolocation). The head was bent ventrally when swimming in such a way that the long tusk was approximately parallel to the axis of the body. The extremely salient occipital condyles of Odobenocetops are indicative of great mobility of the neck, probably related to bottom-feeding. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 134 , 423–452.     

Mitochondrial DNA phylogeny and morphological diversity in the genus Mastus (Beck, 1837): a study in a recent (Holocene) island group (Koufonisi, south-east Crete)

Three endemic Cretan land snail species of the genus Mastus (Beck, 1837) from the island group of Koufonisi (south-east Crete) and the eastern part of Crete, were studied by multivariate analysis of shell morphology and analysis of mtDNA sequences. The phylogeny of the populations studied and the processes effecting the genetic and morphological diversity of the species were investigated. Extremely high mtDNA sequence divergence was observed both within and between populations. The Cretan populations could not be distinguished morphologically, while the populations of the islets were more distinct. We argue that the active geological past of the area (including sea level changes) and the long-term presence of humans has produced a mixing up of Mastus populations leading to the accumulation of high divergence of mtDNA sequences on a small spatial scale. The limited morphological diversity and the distinct shell 'identity' of the islets' populations can be attributed to the selective pressures of the island group.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78, 383–399.