Anuran amphibians have intensively been studied to understand Amazonian biodiversity. Improved methods and sampling has revealed that many widespread nominal species in fact are complexes of species with smaller allopatric ranges. Pan-Amazonian anuran species are rather an exception. In a case study using the three-striped poison frog (Anura: Dendrobatidae: Ameerega trivittata), we ask how the pan-Amazonian distribution of this taxon can be explained and hypothesize that dispersal has played a major role. Species delimitation and intraspecific relationships of the study species were examined from novel and existing (GenBank) sequences of the mitochondrial 16S rRNA gene from 108 specimens of 38 localities using maximum likelihood and Bayesian methods. We performed BioGeoBEARS models using a time-calibrated population tree to reconstruct the biogeographic history. Our results support that A. trivittata is a pan-Amazonian species scattered over its geographic range. Being of Late Miocene origin, the species rapidly spread into newly available space and repeatedly dispersed for-and backward, while vicariance played a major role only in the Early Pliocene. We suggest that intrinsic morphological and life history characteristics (adult size, relative reproductive success) make A. trivittata a more successful disperser than other species, so that riverine barriers are more permeable and hamper allopatric speciation. We conclude that there is no universal causality explaining Amazonia biodiversity, because species-specific biological characteristics are key determents of biogeographical histories. Comparatively better dispersal advantages foster larger geographic ranges and can explain pan-Amazonian distributions.

Abstract Cacti with the common name cholla are classified into several genera of the subfamily Opuntioideae (Cactaceae), and appear to be a monophyletic group. Although recent studies have provided strong resolution of the base of this group, assessment of evolutionary processes within genera have been limited due to taxon sampling. Analysis of four spacer and intron sequences of the chloroplast genome reveals evolutionary patterns and trends that are largely congruent with other recent studies. We concur with recent calls for the recognition of Micropuntia pulchella as monotypic and distinct from Grusonia, the name we use to refer to all species sometimes classified under Marenopuntia and Corynopuntia. Grusonia is monophyletic, and Cylindropuntia is borne on two clades of differing branch lengths. By incorporating the elements of ploidy, reproduction, and sequence divergence, we provide new insight into allopolyploid ancestry, the adaptive impact of ploidal-level variation in species, and the role of reproductive mode in diversification rates in the chollas.

Silene nelsonii M.R. Mesler, M.S. Mayer, and S.K. Carothers (Caryophyllaceae), a species with large, dayblooming flowers from the Trinity River Basin of northwestern California, is described and illustrated. In the past, plants assignable to this new species have been referred to incorrectly as S. bolanderi A.Gray or S. hookeri Nutt. subsp. bolanderi (Gray) Abrams, names that rightfully apply to another member of the Silene hookeri complex. Here we argue that both S. nelsonii and S. bolanderi are worthy of recognition as separate species based on morphological distinction and a molecular phylogenetic analysis. Silene nelsonii differs from S. bolanderi by its more deeply lobed, white petals; lack of well-developed coronal petal appendages; densely ciliate petal bases; and shorter, sometimes branched sepal hairs. We present a key to the five members of the S. hookeri complex, all of which are in some degree rare in California and worthy of conservation concern.

We studied the knowledge gap in GenBank with regard to the ca. 600 anuran species from Amazonia. The markers 12S, 16S, COI and cytb were examined, on which information was available for about half of all species. Both the number of sample sites and the number of samples per species varied greatly (best studied each in 16S: 4.85 +/- 10.37; 11.19 +/- 31.20), and merely one fifth of all species had at least 5 sample sites. This suggests that a considerable portion of species is underrepresented in GenBank. Representativeness is especially difficult to assess in widespread species that at the same time could well represent cryptic allopatric species (i.e., with smaller distributions). This is a well-known phenomenon in Amazonian anurans considering that truly widespread species do exist. Moreover, limited sampling may not necessarily be the result of limited representativeness, as numerous species are known to occupy relatively small localised to regional ranges only. Our study furthermore revealed that in a geographic context, major portions of Amazonia have as yet been undersampled. That is, the total of 453 sample sites (most with more than one species sampled) are spatially clustered, often in areas with increased anthropogenic activity. We conclude that there is a large knowledge gap in terms of spatial sampling, resulting in taxonomic deficiencies.

Although the distribution patterns of major amphibian lineages are mainly explained by a Pangean origin with subsequent vicariant diversification, dispersal events have exerted a strong influence on present-day distributions. Long-distance dispersal (LDD) involves movements outside the standard geographic limits and outside the genetic neighbourhood area of individuals. Although considered ‘rare’, LDD is disproportionately important to amphibian populations, species and communities. To understand the role of LDD in shaping current biogeographic patterns in these tetrapods, we reviewed the cases reported in the literature. A systematic bibliographic search was performed to obtain information on how many studies have dealt with LDD in amphibians, which methods they used, which taxa and distances were involved, and when/where events had apparently occurred. In 41 studies, we recovered at least 90 LDD events (3 active, 87 passive) involving at least 56 extant species and 38 genera. Most events (73) involved the colonization of islands, with rafting being suggested as the most conceivable means of overwater passive dispersal for these vertebrates. In this review, we show that LDD events have played an important role in shaping current amphibian biogeographic patterns, especially the occurrence of disjunct distributions and the colonization of islands.

Neotropical poison frogs (Aromobatidae, Dendrobatidae) are known for their complex behavior including site fidelity and home range maintenance. It has been shown in a few poison frog species that these amphibians are able to return to their home ranges after experimental translocation. In this study we asked if Ameerega trivittata can be allocated to the species performing homing behavior. In this taxon, males and females show home range behavior, while sexes were not distinguished in our study. Fieldwork was carried out in a wild population at Panguana (Peru), using replacement distances of 150 m, 600 m and 900 m. In total, 79 frogs were translocated. Most rapidly returned to their home ranges from all translocation distances, with a decrease of the homing success with longer distance. Among the poison frogs studied so far, it is remarkable that A. trivittata is the only one known to be able to return from 900 m (which perhaps is a remarkable homing distance for anurans in general), while maximum return distances in other species are less than 50% of this. Ameerega trivittata is one of the largest poison frogs (maximum snout-vent length 55 mm). However, long distance homing was not explained by the species' body size. We rather expect that 'good knowledge' of the general area (in terms of integration of learned landmarks) that frogs live in is the reason for the ability of long distance homing in our focal taxon.

Neotropical poison frogs (Dendrobatidae) display complex social behaviors. We studied home range behavior in Ameerega trivittata in the rainy season of two consecutive years at Panguana, Peru. Minimum convex polygon (MCP) and Kernel methods were employed. For the first time in the genus Ameerega, we found that females also occupied home ranges. Home range size in both sexes varied greatly and overlap was commonly found between and between sexes. Only a few specimens (both sexes) were rediscovered in the second rainy season, but always within the home range of the previous year. This finding suggests strong site fidelity but also a high seasonal turnover, which could be an indication of dispersal propensity. Maintenance and size of home ranges in males might be linked to individual fitness and the monopolization of resources (i. e., habitat structures) related to mating. Female home range behavior remains to be explained.

Calls are important premating isolating barriers in frogs; therefore, studying intraspecific variation in calls might allow the assessment of patterns of call divergence during the early stages of speciation. Ranitomeya imitator, a species of dart-poison frog (Dendrobatidae), has undergone extensive color-pattern diversification through a Mullerian mimetic radiation, establishing four distinct morphs in north-central Peru (striped, banded, varadero, and spotted). Partial reproductive isolation exists between certain color morphs, although the specific mechanisms responsible for this isolation are poorly understood. We conducted a species-wide analysis of variation in advertisement calls to investigate whether distinct mimetic morphs show advertisement call differences. We found that different color morphs generally show weak or no differences in advertisement calls, with two exceptions. First, call pulse rates differed between the striped and banded morphs, and that difference coincides geographically with the mimetic transition zone. Second, there is a difference in both note length and dominant frequency between the striped and varadero morphs, which is also geographically coincident with the mimetic transition zone. Future work in this system should attempt to measure the relative importance of different mating cues ( color-pattern, body size, and advertisement calls) mediating mate choice in this species.

We studied territorial interactions with respect to inter-and intraspecific communication of Ranitomeya imitator with the syntopic species Ranitomeya variabilis. These poison frogs (Dendrobatidae) are phenotypically very similar and are considered to be Mullerian mimics. Males of both species use calls for intraspecific communication, but only R. imitator males use calls to advertise and defend their territories. We hypothesized that, to avoid the risk of 'wasting' energy for territorial defence, R. imitator males should be able to discriminate between conspecifics and its model species R. variabilis by the structure of its call. We conducted phonotaxis field experiments attracting wild R. imitator males towards (1) an R. imitator advertisement call, (2) an R. variabilis call, (3) four modified calls with the temporal call parameters note length and internote interval substituted between the two species, and (4) two R. imitator calls at different dominant frequencies beyond the call's known natural range. Male R. imitator did not show responses towards any R. variabilis call (original or modified to match the R. imitator call), but did approach the sound source during playbacks of R. imitator calls with properties of R. variabilis. They also approached their own advertisement call when the dominant frequency was modified. This shows that modified call parameters did not have significant effects on R. imitator males' ability to discriminate their own call from that of R. variabilis. We conclude that the call parameters note length, internote interval or dominant frequency are not essential for call discrimination in R. imitator, perhaps suggesting that among Mullerian mimic anurans, discrimination of conspecifics and heterospecifics acts at a more complex level. (C) 2013 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

The Mason Valley cholla, Cylindropuntia ×fosbergii (C. B. Wolf) Rebman, M.A. Baker & Pinkava, is the putative hybrid of C. bigelovii (Engelm.) F. M. Knuth and some other species of Cylindropuntia. We used AFLPs to screen chollas of the Anza-Borrego Desert in southern California to test this hypothesis of hybrid origin and identify the parental species involved. Other species scrutinized as potential parents include C. echinocarpa (Engelm. & J. M. Bigelow) F. M. Knuth, C. ganderi (C. B. Wolf) Rebman & Pinkava, C. californica var. parkeri (J. M. Coulter) Pinkava, and C. wolfii (L. D. Benson) M.A. Baker. Patterns of band sharing clearly testify to the close relationship between C. ×fosbergii and C. bigelovii. None of the other species screened came close to that level of similarity. Moreover, the numbers of total loci and unique loci in C. ×fosbergii do not meet the expectations of a hybrid taxon. We propose the alternative hypothesis that C. ×fosbergii is the sister species of C. bigelovii.

Previous studies have postulated that the Streptanthus glandulosus Hook. complex (Brassicaceae) originated through gradual range contraction and isolation of populations of an ancestral species distributed in California, United States. Here, we integrate patterns of interfertility, morphology, ITS, and chloroplast DNA (cpDNA) variation to generate a classification system reflective of the phylogeny of this group. As a result, we propose a single species, S. glandulosus, with ID subspecies, to constitute the complex. Populations of S. glandulosus subsp. glandulosus distributed north of the San Francisco Bay are now divided between two new subspecies, S. glandulosus subsp. arkii M. S. Mayer and subsp. raichei M. S. Mayer, diagnosable through molecular apomorphies, perianth color, and fruit orientation. The name S. peramoenus Greene is synonymized to S. glandulasus subsp. glandulasus and is further lectotypified.

Sequence data from the nuclear ribosomal internal transcribed spacer (ITS) region and plastid gene ndhF of 95 species, represented by 147 accessions, were used to determine the tribal limits, monophyly status, and phylogenetic intra-tribal relationships of genera within the New World tribe Schizopetaleae (formerly Thelypodieae; Brassicaceae). Maximum parsimony, Bayesian, and maximum-likelihood analyses all support the separate recognition of three distinct and generally well-supported clades. The largest corresponds to the North–South American Thelypodieae (27 genera: Catadysia , Caulanthus , Chaunanthus , Chilocardamum , Chlorocrambe , Coelophragmus , Dictyophragmus , Dryopetalon , Englerocharis , Hesperidanthus , Mostacillastrum , Neuontobotrys , Polypsecadium , Pringlea , Pterygiosperma , Romanschulzia , Sibara , Sibaropsis , Stanleya , Streptanthella , Streptanthus , Thelypodiopsis , Thelypodium , Thysanocarpus , Warea , Weberbauera , and Werdermannia ). The remaining six South American Schizopetaleae genera were divided into two clades: SCHIZ I of four genera ( Aschersoniodoxa , Brayopsis , Eudema , and Onuris ) and SCHIZ II Schizopetaleae s. str.: Mathewsia , and Schizopetalon ). Resolution within the Thelypodieae clade was limited in both the ITS- and ndhF-based phylogenies. Based on ITS sequence data, elements of the Old World tribe Brassiceae were sister to the Thelypodieae, whereas the ndhF data strongly supported the Sisymbrieae as sister to the Thelypodieae, and the Brassiceae as sister to both tribes. Sister groups to the Schizopetaleae clades I and II were not clearly resolved. Morphological and cytological data support the separation of these three clades. In SCHIZ II, the sepals are always erect to form a closed tube, and the trichomes are mostly dendritic, although other types also occur. In both the Thelypodieae and SCHIZ I clades, the sepals are variously oriented but never form a closed tube, whereas the trichomes are either absent, or simple, forked, or rarely dendritic. The latter clade differs from both SCHIZ II and Thelypodieae by lacking trichomes or rarely having forked ones and by having exclusively basal, entire leaves. Thelypodieae circumscription should be restricted in future to the Thelypodieae clade, Schizopetaleae circumscription to the SCHIZ II clade, while a new tribe is needed for the SCHIZ I clade.

Streptanthus glandulosus Hooker subsp. josephinensis Al-Shehbaz & M. Mayer (Brassicaceae) is described as new. The new combinations S. glandulosus subsp. niger (Greene) Al-Shehbaz, M. Mayer & 1). W. Tavlor, S. glandulosus subsp. sonomensis (Kruckeberg) M. Mayer & 1). W. Taylor, and S. vimineus (Greene) Al-Shehbaz & D. W. Taylor, as well as a new stains for S. glandulosus subsp. albidus (Greene) Al-Shehbaz, M. Mayer & 1). W. Taylor and S. glandulous subsp. hoffmanii (Kruckeberg) M. Mayer & D. W. Taylor, are proposed. A key to the eight subspecies of S. glandulosus is presented. Mesoreanthus fallax Greene and S. sparsiflorus Rollins are reduced to synonymy of S. vimineus and S. platycarpus A. Gray, respectively. The names M. vimineus Greene and S. niger Greene are lectotypified. The IUCN status of S. glandluosus subsp. josephinensis is given as Critically Endangered (CE).

The Brassicaceae is a large plant family (338 genera and 3,700 species) of major scientific and economic importance. The taxonomy of this group has been plagued by convergent evolution in nearly every morphological feature used to define tribes and genera. Phylogenetic analysis of 746 nrDNA internal transcribed spacer (ITS) sequences, representing 24 of the 25 currently recognized tribes, 146 genera, and 461 species of Brassicaceae, produced the most comprehensive, single-locus-based phylogenetic analysis of the family published to date. Novel approaches to nrDNA ITS analysis and extensive taxonomic sampling offered a test of monophyly for a large complement of the currently recognized tribes and genera of Brassicaceae. In the most comprehensive analysis, tribes Alysseae, Anchonieae plus Hesperideae, Boechereae, Cardamineae, Eutremeae, Halimolobeae, Iberideae, Noccaeeae, Physarieae, Schizopetaleae, Smelowskieae, and Thlaspideae were all monophyletic. Several broadly defined genera (e.g., Draba and Smelowskia) were supported as monophyletic, whereas others (e.g., Sisymbrium and Alyssum) were clearly polyphyletic. Analyses of ITS data identified several problematic sequences attributable to errors in sample identification or database submission. Results from parsimony ratchet and Bayesian analyses recovered little support for the backbone of the phylogeny, suggesting that many lineages of Brassicaceae have undergone rapid radiations that may ultimately be difficult to resolve with any single locus. However, the development of a preliminary supermatrix including the combination of 10 loci for 65 species provides an initial estimate of intertribal relations and suggests that broad application of such a method will provide greater understanding of relationships in the family.

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. Lens includes L. culinaris subsp. culinaris (the cultivated lentil) and several wild species distributed from the Mediterranean region to western Asia. We compared sequence variation in the ITS region among species of Lens in an effort to end persisting uncertainty regarding the phylogeny of the genus. The parsimony analysis revealed a single minimum-length tree with a topology congruent with patterns derived by previous studies of nuclear and chloroplast DNA RFLPs. The basal and highly divergent status of the L. nigricans clade is depicted, and the progenitor-derivative relationship between L. culinaris subsp. orientalis and L. culinaris subsp. culinaris is reaffirmed. Resolution in the tree was improved by combining the ITS data set with a pre-existing set of chloroplast DNA restriction site data obtained from the same group of samples.

Morphology of cysts of fairy shrimp (Crustacea, Anostraca) is often very usem in distinguishing between genera and some morphologically distinct species. However, the cysts of many species are indistinguishable from one another due to high interspeclfic morphological similarity or to extensive intra-qxdic variability. In addition, hybridization between some species is possible and unlikely to be detected by examining cyst morphology. We have developed an approach basedon RAPD-PCR molecular markers that allows for the rapid and accurate identification of fairy shrimp species at the cyst stage. This technique can quickly differentiate between three congenic species of potentially c o a g fa j r shrimp in southern California that are known to hybridize and whose cysts are dif3cult or impossible to identify using morphological characters. This technique has the potential to be expanded to incorporate many more species. Key word!: California, cysts, branchinectids, Brachiopoda, ephemeral wetlands, fajr shnmp, RAPD-PCR analysis 2001 TRANSACnONS OF THE WESTERN SECnON OF THE WlLDLlFE SOCI€TY37:1621 California's array of ephemeral wetlands (vernal pools, playas, etc.) provide habitat for diverse and largely endemic flora and fauna (Holland and Jain 1988, Simovich 1998). The loss of ephemeral pools in California has caused numerous species of plants and animals associated with these habitats to become federally listed as threatened or endangered, including several species of fairy shnmp (CNstacea: Anostraa) (Federal Register 1988; 1993; 1994a, b; 1997). C o n s e q u e , docmentingthe distribution of endangered species, compiling environmental impact reports, and devising consemation and management plans has made accurate identification of species increasingly important. Two sensitive ephemeral pool animals are the endemic Californian fairy shrimp species, Branchinecta sandiegonensis (Fugate 1993) and B. lynchi (Eng et al. 1990). BcRh of these two species may b e in the same ephemeral pools or pools in the same area as the more common and unthreatened species B. linhhli (Eng et al. 1990, Fugate 1992, Simovich and Fugate 1992, Eriksen and Bek 1999). Branchinecta sandiegonensis and B. lynchi are endangered and threatened, respecbvely, and are both subjects of consemation efforts. Hybridization involving rare species is considered a threat to the persistence of those species for a myriad of reasons, including genetic swamping, outampetition, andincreased disease sensitivity (Levln et al. 19%). All three possible combinations of southern Californian branchinectid hybrids have been produced in the laboratory (Fugate 1992). Although no hybrids have yet been observed in natural populations, the possibility of their existence is a conservation concern. The ephemeral wetlands in which these anostracans are found are characterized by periodic hydration and dehydration events. Hydrations may occur years apart and may last from days to months. Due to the short duration and irregular periodicity of this habitat, the adult forms may be present for only a fav weeks out of the year, if at all. Thus, the population usually exists predominately in the soil as encysted embryos. Identification of cysts of genus Branchinecta is possible by examining their morphology, but species identification is oftenproblematic (Mura 1991a, Hill and Shepard 1997). Previously, the only way to positively identify most branchinectid cysts to the level of species has been to hyhtethecysts, reartheduhp, andihtiQtheaWts based on morphological characteristics (see Bek 1975, Eriksen and Be& 1999). Unfortunately, this ppnx;ess may take weeks to complete. Furthermore, hydration conditions must be suitable to all species present or some may not hatch or survive to adulthood and possibly go undetected. Efforts have also been made to distinguish the cysts of anostracans using scanning electron microscopy (SEM). Cyst diameter has been abandoned as a diagnostic character due to great variability within species (Be& et al. 1990, Mura 199 lb) and most attention has focused on detecting species differences in tertiary envelope a p 'Address all correspondence to this author. Ernail: simo@sandiego.edu 16 TRANS.WEST.SECT.WILDL.SOC. 37:2001 Identification of Branchinectid Cysts 0 Moorad, et al. 17 pearance (Mum and Thiery 1986; Mum 199 la, b, c; Hill and Shepard 1997). Unfortunately, this approach requires dZ€icult and often arguably subjective judgment regarding morphology. In addition, tertiary envelope characteristics can be affected by non-genetic factors such as population elevation, maternal stress, preservation conclitim, cyst age, and even SEM sample preparation (Mura 1991~). These fixtors, coupled with high within-species mrphological variabilrty (Mura 199 la, c; Hill and Shepard 1997), compromise a>nfin be flexlble enough to allow for large amounts of within-species variability; offer enough resolving power to distinguish not only between species but hybrids of closely related species; and examine characters that are immune to environmental modification Genetic markers have proven useful in identilication efforts for diverse systems at morphologiicallysimilartaxa(forreviewseeHadrysetal. 1992). For example, biochemical markers have also been used to investigate phylogenetics and population genetics of branchinectids (Fugate 1992, Davies et al. l997, Bohonak 1998) but samples have been restricted to adult tissues. This approach works well when dealing with adult forms or otherwise large organisms but it carries substantial tissue requirements. Because branchinectid cysts are several orders of magnitude smaller than the adult forms (4-5 pg each), analysis by either of these techniques is not possible. The polymerase chain reaction (FTR) offers a solution to the minimal tissue problem. When used in conjunction with randomly amplifiedpolymorphic DNA (MPD) primers, anonymous DNA-ents (or markers) ofdifferent sizes can be amplified and visualized on an agarose gel (Welsh and McClelland 1990, W* et al. 1990). This technique can provide access to a very large number of genetic markers. These markers can resolve genetic differences between individuals, populations, species, and genera (for raiew see Hadrys et al. 1992). Recently, RAPD markers have been shown to be very useful in species and subspecies identification of small invertebrates (Ballinger-Crabtree et al. 1992, Coffroth and Mulawka 1995, Humbert and Cabaret 1995). The purpose of this study was to devise an approach for the rapid and accurate identification of cysts of morphologically similar southern Californian branchinectid species (B. Table 1. Sample locations in California for populations of 3 species of fairy shrimp (Branchinecta sp.) species Localtty Collectors B. sandiegonensis 1. Nobel Drive, San Diego County 2. Otay Mesa, San Diego County 3. Miramar NAS, San Diego County B. lynchi 4. Santa Rosa Plateau Reserve, Riverside County 5. Corning, Tehama County 6. Truckee Creek, Tehama County 7. Vim, Tehama County