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new & recent described Flora & Fauna species from all over the World esp. Asia, Oriental, Indomalayan & Malesiana region
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    Rodriguezia adani
    Alvarez & Villalobos, 2018


    A new species of freshwater crab of the family Trichodactylidae, genus Rodriguezia Bott, 1969 is described from Grutas de Agua Blanca in southern Tabasco, Mexico. Rodriguezia is a genus endemic to northern Chiapas and southern Tabasco, distributed over a small area of 70 km.Rodriguezia adani n. sp., the third species of the genus, occurs north of its two congeners, being stygobitic with obvious adaptations to cave life. It can be distinguished from R. villalobosi, an epigean species, by the absence of eyes, lack of pigmentation and elongation of the pereiopods; and from R. mensabak by having less elongated pereiopods relative to carapace breadth, an extremely reduced ocular peduncle, and a smaller adult size.

    Keywords: Crustacea, Trichodactylinae, stygobitic, Grutas de Agua Blanca, Tabasco, Chiapas

    FIGURE 2. Rodriguezia adani n. sp. male holotype: dorsal view. 

    Rodriguezia adani n. sp.

    Distribution. The new species is only known from Grutas de Agua Blanca, Macuspana, Tabasco, Mexico.

    Etymology. We name the new species after Adán Gómez-González, explorer, biologist and friend, who found these crabs while exploring caves in Tabasco and Chiapas, Mexico.

    Fernando Alvarez and José Luis Villalobos. 2018. A New Species of Stygobitic Freshwater Crab of the Genus Rodriguezia Bott, 1969 (Crustacea: Decapoda: Trichodactylidae) from Tabasco, Mexico.  Zootaxa. 4378(1); 137-143. DOI:  10.11646/zootaxa.4378.1.10

    Dedican Nueva Especie de Crustáceo al Joven Biólogo Asesinado en Chiapas:"Rodriguezia adani"... - Biosfera 10 via @@biosferadiez

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    Hyperolius stictus 
    Conradie, Verburgt, Portik, Ohler, Bwong & Lawson, 2018


    A new species of African reed frog (genus Hyperolius Rapp, 1842) is described from the Coastal Forests of the Eastern Africa Biodiversity Hotspot in northeastern Mozambique. It is currently only known from less than ten localities associated with the Mozambican coastal pans system, but may also occur in the southeastern corner of Tanzania. Phylogenetic reconstructions using the mitochondrial 16S marker revealed that it is the sister taxon of Hyperolius mitchelli (>5.6% 16S mtDNA sequence divergence) and forms part of a larger H. mitchelli complex with H. mitchelli and H. rubrovermiculatus. The new species is distinguished from other closely related Hyperolius species by genetic divergence, morphology, vocalisation, and dorsal colouration.

    Keywords: Amphibia, Amphibian, endemic, coastal pans

     Werner Conradie, Luke Verburgt, Daniel M. Portik, Annemarie Ohler, Beryl A. Bwong and Lucinda P. Lawson. 2018. A New Reed Frog (Hyperoliidae: Hyperolius) from coastal northeastern Mozambique. Zootaxa. 4379(2); 177–198.   DOI:  10.11646/zootaxa.4379.2.2

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    A herd of Saurolophus angustirostris moves along a river bank after a storm in the Cretaceous Nemegt Basin. The feet of the large herbivores sink into the soft sediment crushing the skull of a Tarbosaurus bataar that was lying in the mud.

     Illustration based on specimen MPC-D107/05 collected at the Nemegt locality (Nemegt Formation) and discovered by J.Ed. Horton. Artwork by Davide Bonadonna.

    Fanti, Bell, Currie & Tsogtbaatar, 2018. 
      Palaeogeography, Palaeoclimatology, Palaeoecology. 494

    • The Nemegt Basin is perhaps the most important fossil-bearing region of Mongolia.
    • The unique fossils of Mongolia have sparked an explosion of illegal fossil poaching in the country.
    • We introduce multidisciplinary methodologies to understand the Cretaceous Nemegt ecosystem.
    • We discuss biotic response to local and large-scale Nemegt paleocological dynamics.

     Keywords: Mongolia, Late Cretaceous, Paleoecology, Stratigraphy, Vertebrate paleontology

    Fig. 1: A herd of Saurolophus angustirostris moves along a river bank after a storm in the Cretaceous Nemegt Basin. The feet of the large herbivores sink into the soft sediment crushing the skull of a Tarbosaurus bataar that was lying in the mud. Illustration based on specimen MPC-D107/05 collected at the Nemegt locality (Nemegt Formation) and discovered by J.Ed. Horton.
    Artwork by Davide Bonadonna. 

      Federico Fanti, Phil R. Bell, Philip J. Currie and Khishigjav Tsogtbaatar. 2018. The Nemegt Basin — One of the Best Field Laboratories for Interpreting Late Cretaceous Terrestrial Ecosystems [Dedicated to Ryszard Gradziński, Ivan Antonovĭc Efremov, and Demchig Badamgarav whose pioneer work unraveled the unique Late Cretaceous Nemegt ecosystems.]. [in Federico Fanti, Phil Bell, Philip Currie and Khishigjav Tsogtbaatar (eds.). 2018. The Late Cretaceous Nemegt Ecosystem: Diversity, Ecology, and Geological Signature.Palaeogeography, Palaeoclimatology, Palaeoecology. 494; 1-4. DOI: 10.1016/j.palaeo.2017.07.014

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    Sauripes hadongensis Lee, Lee, Fiorillo & Lü, 2018

    A reconstruction of a lizard running bipedally chased by the pterosaur 
    Pteraichnus koreanensis, based on the trackway.  
    Illustration: Chuang Zhao

    Four heteropod lizard trackways discovered in the Hasandong Formation (Aptian-early Albian), South Korea assigned toSauripeshadongensis, n. ichnogen., n. ichnosp., which represents the oldest lizard tracks in the world. Most tracks are pes tracks (N = 25) that are very small, average 22.29 mm long and 12.46 mm wide. The pes tracks show “typical” lizard morphology as having curved digit imprints that progressively increase in length from digits I to IV, a smaller digit V that is separated from the other digits by a large interdigital angle. The manus track is 19.18 mm long and 19.23 mm wide, and shows a different morphology from the pes. The predominant pes tracks, the long stride length of pes, narrow trackway width, digitigrade manus and pes prints, and anteriorly oriented long axis of the fourth pedal digit indicate that these trackways were made by lizards running bipedally, suggesting that bipedality was possible early in lizard evolution.

    Figure 1 Photograph and drawing of lizard trackways on the block.

    Figure 5 A reconstruction of a lizard running bipedally chased by the pterosaur Pteraichnus koreanensis, based on the trackway (Drawn by Chuang Zhao).

    Systematic ichnology
    Order Squamata Oppel, 1811

    Sauripeshadongensis ichnogen. et ichnosp. nov.

    Etymology: Ichnogenus named from ancient Greek “sauros” (lizard) and “pes” (foot). Ichnospecies named after Hadong County that yielded the holotype.

    Holotype: Manus and pes prints on a mudstone slab (70 × 30 cm) (KIGAM VP 201501: Korea Institute of Geoscience and Mineral Resources, Vertebrate Paleontology).

    Type locality and horizon: Hasandong Formation, Lower Cretaceous (Aptian-early Albian), an abandoned quarry next to Hadong power plant, Hadong County, South Gyeongsang Province, South Korea.

    Diagnosis: Quadrupedal tracks; manus prints are medial to the pes prints; the pes prints are larger than the manus prints; plantigrade and pentadactyl pes prints are longer than wide; the digit length progressively increasing from digits I to IV (ectaxonic); digit V is oriented more laterally and offset from other digits; digit imprint IV is more than twice the length of the metatarsal impression; plantigrade and pentadactyl manus print has similar length and width dimensions; digits II and IV are shorter than digit III (mesaxonic); the interdigital angle between digits I and V of the manus is larger than that of the pes.

    Figure 2 Manus and pes tracks ofSauripes hadongensis, n. ichnogen., n. ichnosp. (a) Enlarged photograph and drawing of a manus imprint (B1). (b) A pes imprint (A6). Scale bars equal 1 cm.

    Figure 3 Pes tracks of Sauripes hadongensis, n. ichnogen., n. ichnosp. (a) Enlarged photograph and drawing of a pes imprint (A3). (b) A pes imprint (B8). (c) A pes imprint (B9). Scale bars equal 1 cm.

    Hang-Jae Lee, Yuong-Nam Lee, Anthony R. Fiorillo and Junchang Lü. 2018. Lizards ran Bipedally 110 Million Years Ago.  Scientific Reports. 8, Article number: 2617.  DOI: 10.1038/s41598-018-20809-z

    Fossil Footprints Are Oldest Traces of Lizards Running on Two Legs via @NatGeo

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    Callipia rosetta Thierry-Mieg, 1904
    C. walterfriedlii  Brehm, 2018
    C. augustae Brehm, 2018

       DOI:  10.5852/ejt.2018.404 


    The vividly coloured Neotropical genus Callipia Guenée (1858) (Lepidoptera Linnaeus, 1758, Geometridae (Leach, 1815), Larentiinae (Leach, 1815), Stamnodini Forbes, 1948) is revised and separated into four species groups, according to a provisional phylogeny based on Cytochrome Oxidase I (COI) gene data and morphology. 

    Fourteen new species are described using COI data and morphology:
    a) in the balteata group: C. fiedleri sp. nov.,C. jakobi sp. nov., C. lamasi sp. nov.;
    b) in the vicinaria group:C. hausmanni sp. nov., C. walterfriedlii sp. nov.;
    c) in the parrhasiata group: C. augustae sp. nov.,C. jonai sp. nov., C. karsholti sp. nov.,C. levequei sp. nov., C. milleri sp. nov., C. sihvoneni sp. nov., C. wojtusiaki sp. nov. and
    d) in the constantinaria group: C. hiltae sp. nov., C. rougeriei sp. nov.
     One new subspecies is described: C. wojtusiaki septentrionalis subsp. nov. 

    Two species are revived from synonymy:C. intermedia Dognin, 1914 stat. rev. and C. occulta Warren, 1904 stat. rev. 

    The taxon hamaria Sperry, 1951 is transferred from being a junior synonym of C. constantinaria Oberthür, 1881 to being a junior synonym of C. occultastat. rev. The taxon admirabilis Warren, 1904 is confirmed as being a junior synonym of C. paradisea Thierry-Mieg, 1904. The taxon languescens Warren, 1904 is confirmed as being a junior synonym of C. rosetta, Thierry-Mieg, 1904 and the taxon confluens Warren, 1905 is confirmed as being a junior synonym of C. balteata Warren, 1905. 

    The status of the remaining species is not changed: C. aurata Warren, 1904, C. brenemanae Sperry, 1951, C. parrhasiata Guenée, 1858, C. flagrans Warren, 1904, C. fulvida Warren, 1907 and C. vicinaria Dognin. 

    All here recognised 26 species are illustrated and the available molecular genetic information of 25 species, including Barcode Index Numbers (BINs) for most of the taxa is provided. The almost threefold increase from 10 to 26 valid species shows that species richness of tropical moths is strongly underestimated even in relatively conspicuous taxa. Callipia occurs from medium to high elevations in wet parts of the tropical and subtropical Andes from Colombia to northern Argentina. The early stages and host plants are still unknown.

    Keywords: Callipia; taxonomy; Andes; insect; Neotropics

    Figs 131–138. Living specimens and habitats. 131. Callipia rosetta Thierry-Mieg, 1904, ♂, Ecuador, Loja province, Podocarpus National Park, Cajanuma, 2897 m, 26 Mar. 2011. The specimen was attracted to light and benumbed. 132. Elfin forests are a habitat of C. rosetta Thierry-Mieg, 1904 and C. walterfriedlii sp. nov., Ecuador, Loja province, Podocarpus National Park, Cajanuma, 3000 m, 30 Jan. 2013. 133. C. walterfriedlii sp. nov., ♀, Ecuador, Loja province, Podocarpus National Park, Cerro Toledo, 2938 m, 27. Feb. 2013. The specimen was attracted to light and benumbed. 134. Habitat (elfin forest) of C. walterfriedlii sp. nov. at Cerro Toledo. 

    Figs 131–138. Living specimens and habitats. 135. Callipia augustae sp. nov., ♂, Peru, Cusco province, Wayqecha station, 2900 m, 26 Aug. 2016. The specimen was collected at night, trapped, photographed and released the next morning. 136. Habitat of C. augustae sp. nov. and Callipia sp. near Wayqecha station. 137. C. augustae sp. nov., ♂, Peru, Cusco province, road Wayqecha–Pillcopata, 2284 m, 23 Aug. 2016. The specimen was attracted to UV light and tried to take up fluid (see proboscis). 138. Callipia sp. at Wayqecha station, 4 Sep. 2016. This specimen was attracted to UV light, but escaped into the vegetation when disturbed.

    Gunnar Brehm. 2018. Revision of the Genus Callipia Guenée, 1858 (Lepidoptera, Geometridae), with the Description of 15 New Taxa. European Journal of Taxonomy. 404; 1–54.   DOI:  10.5852/ejt.2018.404

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    Tugenchromis pickfordi
     Altner, Schliewen, Penk & Reichenbacher, 2017

    The highly diverse tropical freshwater fish family Cichlidae is sparsely represented in the fossil record. Here we describe the new cichlid †Tugenchromis pickfordi, gen. et sp. nov., from the Upper Miocene (9–10 Ma) of central Kenya. The new taxon possesses a unique combination of characters, including six lateral line foramina on the lacrimal, three lateral line segments, cycloid scales, and a low number of vertebrae (29), dorsal fin spines (13), and dorsal soft rays (9). Its lacrimal morphology and tripartite lateral line suggest an affinity with the present-day Lake Tanganyika tribes Ectodini and Limnochromini, and thus with members of the ‘East African Radiation’ among the African cichlids. To further elucidate the relationships of †T. pickfordi, we used a comprehensive comparative data set comprising meristic data from all present-day tribes of the ‘East African Radiation.’ Principal coordinates analyses support links between the fossil and Ectodini + Limnochromini, and additionally with modern Haplochromini. We conclude that †T. pickfordi could be an extinct lineage within the ‘most ancient Tanganyika tribes,’ or a stem lineage of the ‘ancient Tanganyika mouthbrooders.’ A direct relationship to the Haplochromini is unlikely because its members do not exhibit the derived characteristics of the lacrimal as seen in †T. pickfordi. Because Lake Tanganyika is located in the western branch of the East African Rift System, †T. pickfordi from the eastern branch supports the ‘melting-pot Tanganyika hypothesis,’ which posits that the cichlids of modern Lake Tanganyika are derived from riverine lineages that had already diversified prior to the lake formation.

    FIGURE 2. †Tugenchromis pickfordi, gen. et sp. nov. A1–A2, holotype in part (OCO-5-35) and counterpart (OCO-5-22); A3, right lateral view of the specimen (shading refers to ribs from the left side of the specimen);

    Abbreviations: cl, cleithrum; cor, coracoid; ep, epural; hs, hemal spine; hyp, hypural plate; lac, lacrimal; nlc, neurocranial lateral line canal; ns, neural spine; o, otolith; op, operculum; ph, parhypural; pha, pharyngeal teeth; ppc, postcleithrum; ptt, posttemporal; pu, preural centrum; rad, radials; sca, scapula; scl, supracleithrum; sop, suboperculum; us, urostyle; un1, uroneural 1; = , tubular lateral line scale; °, pitted lateral line scale. 

    CICHLIDAE Bonaparte, 1835

    TUGENCHROMIS, nov. gen.

    Generic Diagnosis: Lateral line on the trunk divided into three segments, two of which are posterior lateral lines. One posterior segment positioned ventrally, the other dorsally to the anterior lateral line segment. This is a condition not seen in any other cichlid genus.

    Etymology: Tugen’ refers to the ‘Tugen Hills’ (named after the local people, i.e., the ‘Tugen,’ a subgroup of the Kalenjin ethnic group), in which the type locality of the new fossil taxon is located. The Greek word ‘Chromis’ (χρόμις) is a name used by the Ancient Greek and was applied to various fish. It is a common second element in cichlid genus names. Tugenchromis is masculine.

    Type Species: Tugenchromis pickfordi, sp. nov.


    Holotype: OCO-5-22/35, partially complete skeleton in part and counterpart (Fig. 2A1–A3), approximately 60 mm total length, 33.5 mm body length.

    Etymology: Species named in honor of the paleontologist Martin Pickford in recognition of his outstanding contributions to the geology and paleontology of East Africa.

    Locality, Horizon, and Age: Outcrop Waril in Central Kenya; Ngorora Formation, Member E; late Miocene (9–10 Ma) (see Rasmussen et al., 2017).

    Based on lacrimal morphology and meristic data derived from all present-day cichlids of the ‘East African Radiation,’ we propose that the newly discovered cichlid fossil from the upper Miocene of Central Kenya either represents a stem lineage of the ‘ancient Tanganyika mouthbrooders’ or an extinct lineage within the ‘most ancient Tanganyika tribes.’ This result implies that the use of a comprehensive set of comparative material derived from extant cichlids may make it possible to phylogenetically place other fossil cichlids with greater confidence in future studies.

    Apart from a lower Miocene cichlid from Uganda (‘cf. Pelmatochromis spp.’), none of the previously described fossil cichlid taxa from Africa, Arabia, and Europe possess distinctive similarities to †T. pickfordi. This indicates that the Ngorora fish Lagerstätte in Central Kenya may provide an unrivalled window into the evolutionary history of African cichlids, particularly into the origin of the ‘East African Radiation,’ i.e., the megadiversity of the present-day cichlids in Lake Tanganyika, Lake Malawi, and Lake Victoria.

    Furthermore, the new fossil provides additional support for the presence of an ancient east-west connection (e.g., proto-Malagarasi River) between the Central Kenya Rift and Lake Tanganyika, which is consistent with previous assumptions regarding the hydrological networks across East and Central Africa during the Miocene.

    Melanie Altner, Ulrich K. Schliewen, Stefanie B. R. Penk and Bettina Reichenbacher. 2017 . †Tugenchromis pickfordi, gen. et sp. nov., from the upper Miocene—A Stem-group Cichlid of the ‘East African Radiation’. Journal of Vertebrate Paleontology. 37(2); e1297819. DOI:  10.1080/02724634.2017.1297819

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    Liu, Wang, Ke, et al., 2018. 

    1. With accelerating species introductions in an era of globalization, co-occurring alien species have become increasingly common. Understanding the combined ecological impacts of multiple invaders is not only crucial for wildlife managers attempting to ameliorate biodiversity loss, but also provides key insights into invasion success and species coexistence mechanisms in natural ecosystems. Compared with much attentions given to single-invader impacts, little is known about the impacts of multiple co-occurring invaders.
    2. The American bullfrog (Lithobates catesbeianus Rana catesbeiana) and the red swamp crayfish (Procambarus clarkii) are two aquatic invasive species in many different areas of the globe. They coexist with native anurans in a variety of permanent lentic waters, which provide an ideal model system to explore the combined effects of multiple invaders from different trophic levels on native species.
    3. Based on a global diet analysis covering 34 native and invasive bullfrog populations, and data from 10-year field surveys across 157 water bodies in the Zhoushan Archipelago, China, we observed a reduced impact of bullfrogs on native anurans at high crayfish densities when the two invaders co-occurred.
    4. The global diet analysis showed that crayfish occurrence reduced the number of native anuran prey consumed by bullfrogs in both native and invasive populations. After accounting for pseudoreplication of different observations among water bodies, islands, and survey time, model averaging analyses based on GLMMs showed a negative relationship between bullfrog density and native anuran densities for field observations of invasive bullfrogs alone and co-invaded observations with low crayfish density. However, this negative relationship disappeared when the two invaders co-occurred with high crayfish density. Structural equation modelling (SEM) analyses further validated that the impacts of bullfrogs on native frogs were mitigated by the negative interactions between crayfish and bullfrogs.
    5. Our results provide novel evidence of a density-dependent antagonistic effect of two sympatric invaders from different trophic levels on native species. This study highlights the importance of considering complex interactions among co-invaders and native species when prioritizing conservation and management actions and will facilitate the development of a more precise framework to predict invasion impacts.

      Xuan Liu, Supen Wang, Zunwei Ke, Chaoyuan Cheng, Yihua Wang, Fang Zhang, Feng Xu, Xianping Li, Xu Gao, Changnan Jin, Wei Zhu, Shaofei Yan and Yiming Li. 2018. More Invaders Do Not Result in Heavier Impacts: The Effects of Non-native Bullfrogs on Native Anurans are Mitigated by High Densities of Non-native Crayfish.  Journal of Animal Ecology. DOI: 10.1111/1365-2656.12793   

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    Idiosepius hallami
    Reid & Strugnell, 2018


    A new species of pygmy squid, Idiosepius hallami n. sp., is described from eastern Australia. It differs from I. notoides Berry, 1921 and I. pygmaeus Steenstrup, 1881 (also found in Australian waters) in a number of traits, including the number of club suckers, shape of the funnel-mantle locking apparatus and the modification of the male hectocotylus. Mitochondrial DNA markers (12S rRNA, 16S rRNA and cytochrome c oxidase subunit 1) indicate that it is also distinct on a molecular level. The new Australian species is also recognised as the taxon from Stradbroke I., Queensland for which the entire mitochondrial genome has been sequenced (Hall et al. 2014). Idiosepius hallami n. sp. is compared with all nominal Idiosepius Steenstrup, 1881 and a current summary of Idiosepius systematics is provided as a basis for future studies. Based on our analyses, we propose the elevation of the ‘notoides’ clade to the new genus Xipholeptos n. gen., retaining Idiosepius as the genetic epithet for all other nominal idiosepiids. This is supported by: monophyly of the two lineages based on molecular data sets, the level of sequence divergence between these lineages, and morphological differences. The ‘notoides’ clade is endemic to southern Australia and its basal phylogenetic position suggests that the family may have originated in the Australasian region. Idiosepiids are found in seagrass beds and among mangroves—among the most threatened ecosystems in the world.

    Keywords: Mollusca, new taxa, pygmy squid, XipholeptosIdiosepiusIdiosepius hallami, seagrass, Australia

    Idiosepius hallami, attached to a seagrass blade, Cudgen Creek, northern NSW.
     photo: M. Reid

    Xipholeptos notoides n. gen. b, live animal, anterio-lateral view, Victoria, Port Phillip Bay, Point Cooke, photo J. Gaskell. c, live animal, dorsal view, Victoria, Port Phillip Bay, Ricketts Point, photo J. Gaskell. d, e, live animal, ventral view, female, AM C.532745, photos A. Reid. 

    FIGURE 10. Idiosepius hallami n. sp.a, live animal, dorsal view, NSW, Lord Howe Island, photo A. Reid.
    Xipholeptos notoides n. gen. b, live animal, anterio-lateral view, Victoria, Port Phillip Bay, Point Cooke, photo J. Gaskell. c, live animal, dorsal view, Victoria, Port Phillip Bay, Ricketts Point, photo J. Gaskell. d, e, live animal, ventral view, female, AM C.532745, photos A. Reid. 

    Amanda L. Reid and Jan M. Strugnell. 2018. A New Pygmy Squid, Idiosepius hallami n. sp. (Cephalopoda: Idiosepiidae) from eastern Australia and Elevation of the southern Endemic ‘notoides’ Clade to A New Genus, Xipholeptos n. gen.   Zootaxa.  4369(4); 451–486. DOI: 10.11646/zootaxa.4369.4.1

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     Euryrhynchus amazoniensis  Tiefenbacher, 1978

    in Pachelle & Tavares. 2018. 


    The present revision is based on the largest sample of Euryrhynchidae Holthuis, 1950 studied to date, with special reference to Euryrhynchus Miers, 1878. The revision confirms the validity of the 8 currently recognized species of Euryrhynchidae and describes 2 new species related to Euryrhynchus amazoniensis Tiefenbacher, 1978: E. taruman sp. nov. and E. tuyuka sp. nov. The species Euryrhynchus amazoniensis, E. burchelli Calman, 1907, E. pemoni Pereira, 1985 and E. wrzesniowskii Miers, 1878 are redescribed and illustrated based on specimens from the type series and additional material. Additional diagnostic characters are proposed to differentiate the species of Euryrhynchus, previously separated only by the armature of the second pereopod carpus and merus.

    Keywords: Crustacea, Amazon, South America, West Africa, Gondwana, new species

    Paulo P. G. Pachelle and Marcos Tavares. 2018. The Freshwater Shrimp Family Euryrhynchidae Holthuis, 1950 (Crustacea: Decapoda: Caridea) Revisited, with A Taxonomic Revision of the Genus Euryrhynchus Miers, 1878.  Zootaxa. 4380(1); 1-110.   DOI:  10.11646/zootaxa.4380.1.1

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    Lecanorchis sarawakensis Suetsugu & Naiki

    in  Suetsugu,Ling,Naiki,et al., 2018.

    Lecanorchis Blume (1856: 188) comprises about 30 species of mycoheterotrophic orchids (Seidenfaden 1978, Hashimoto 1990, Szlachetko & Mytnik 2000, Govaerts et al. 2017) characterized by having numerous, long, thick, horizontal roots produced from a short rhizome, presence of a calyculus (i.e. a cup-like structure located between the base of the perianth and apex of the ovary) and an elongate column with a pair of small wings on each side of the anther (Seidenfaden 1978, Hashimoto 1990). The genus is distributed across a wide area including China, Korea, India, Indonesia, Japan, Laos, Malaysia, New Guinea, Pacific islands, the Philippines, Taiwan, Thailand and Vietnam (Seidenfaden 1978, Hashimoto 1990, Pearce & Cribb 1999, Szlachetko & Mytnik 2000, Averyanov 2011, 2013).

    Lecanorchis sarawakensis in the type locality.
    A. Habit. B. Flower, side view. C. Flower, front view.

    Lecanorchis sarawakensis Suetsugu & Naiki, sp. nov.

    Kenji Suetsugu,Ling Chea Yiing,Akiyo ,Shuichiro Tagane,Yayoi Takeuchi,Hironori Toyama and Tetsukazu Yahara. 2018. Lecanorchis sarawakensis (Orchidaceae, Vanilloideae), A New Mycoheterotrophic Species from Sarawak, Borneo. Phytotaxa. 388(1); 135–139. DOI:  10.11646/phytotaxa.338.1.13

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    Edmontosaurus annectens (Marsh, 1892)

    in Wosik, Goodwin & Evans, 2018
       DOI: 10.1080/02724634.2017.1398168    

    The Hell Creek Formation preserves one of the most intensely studied late Cretaceous terrestrial fossil units. Over 22 dinosaur genera are currently recognized from this unit, but the record of juvenile individuals is surprisingly limited. Here, we document a nestling hadrosaur that represents the first occurrence of an articulated nestling dinosaur skeleton from the latest Cretaceous (late Maastrichtian) of North America. The specimen (UCMP 128181) preserves a partial scapula, nearly complete rib cage, vertebral series from the shoulder to mid-tail, a large portion of the pelvic girdle, and both hind limbs through a combination of bone and/or natural impressions in the concretion. It is assignable to the genus Edmontosaurus based on the shape of the prepubic process, or blade, of the pubis. The specimen represents the earliest ontogenetic growth stage of Edmontosaurus cf. annectens and possesses a femur length of 148 mm. It greatly contributes as a new end member to a sample of associated Edmontosaurus skeletons that is well suited for allometrically testing the hypothesized ontogenetic gait shift in hadrosaurs from bipedal juveniles to quadrupedal adults using individual limb proportions. Although UCMP 128181 does not preserve forelimbs, regressions based on associated Edmontosaurus skeletons (N = 25) reveal overall isometry of the forelimb relative to the hind limb, and within each limb. These data indicate that Edmontosaurus nestlings were anatomically capable of fully quadrupedal locomotion and provide no compelling evidence to support an ontogenetic gait shift in hadrosaurids.

    UCMP-128181, Edmontosaurus cf. annectens

    Mateusz Wosik, Mark B. Goodwin and David C. Evans. 2018. A Nestling-sized Skeleton of Edmontosaurus (Ornithischia, Hadrosauridae) from the Hell Creek Formation of northeastern Montana, U.S.A., with An Analysis of Ontogenetic Limb Allometry.   Journal of Vertebrate Paleontology.  DOI: 10.1080/02724634.2017.1398168   

    First baby #dinosaur skeleton from the Hell Creek Formation
    Published in @SVP_vertpaleo with Mark Goodwin and @DavidEvans_ROM
    Cover art by @SaurianGame. Funded by @ucmpberkeley Welles Fund.


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    Hyloscirtus japreria 
    Rojas-Runjaic, Infante-Rivero, Salerno & Meza-Joya, 2018


    A new species of Hyloscirtus, belonging to the H. bogotensis species Group, is described from the Venezuelan and Colombian slopes of the Sierra de Perijá. The new species can be readily distinguished from its congeners by the combination of the following characters: mental gland present, disc-shaped and small; ulnar, outer, and inner tarsal folds present; calcar tubercle absent; whitish stripes on external border of upper eyelids and supratympanic folds, longitudinally on the mid-dorsum, on supracloacal fold, outer ulnar folds, inner and outer tarsal folds, and also on dorsal internal surface of shanks. We estimate phylogenetic relationships based on mtDNA (spanning fragments of 12S rRNA, tRNA-Val and 16S rRNA), of all Hyloscirtus species available in Genbank, as well as the new species described herein, H. callipeza, H. jahni, and H. platydactylus, all of which have not been previously sequenced. Our molecular data support the hypothesis of the new species as sister species of H. callipeza and indicates that H. jahni does not belong to the H. bogotensis species Group, but rather is sister species of all other Hyloscirtus (sensu Faivovich et al. 2005). Based on this last result we propose a new species group for H. jahni and the synonymy of Colomascirtus in Hyloscirtus. We also provide the first description of the advertisement call of H. callipeza. With the new species described herein, the number of Hyloscirtus species increases to 37.

    Keywords: Amphibia, Advertisement call, Amphibia, Andes, Colomascirtus, Hylinae, Hyloscirtus bogotensis species Group, Hyloscirtus callipeza, integrative taxonomy, phylogeny

    Fernando J.M. Rojas-Runjaic, Edwin E. Infante-Rivero, Patricia E. Salerno and Fabio Leonardo Meza-Joya. 2018. A New Species of Hyloscirtus (Anura, Hylidae) from the Colombian and Venezuelan Slopes of Sierra de Perijá, and the Phylogenetic Position of Hyloscirtus jahni (Rivero, 1961)Zootaxa. 4382(1);  121–146.  DOI: 10.11646/zootaxa.4382.1.4

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    Australocarcinus insperatus 
    Ng & Daniels, 2018 

    A new species of freshwater chasmocarcinid crab, Australocarcinus insperatus sp. n., is described from the Seychelles Islands in the Indian Ocean. This is the first record of the genus and the subfamily Trogloplacinae Guinot, 1986, from the Indian Ocean, with all other members previously recorded from Australia, New Britain, New Caledonia, and Palau in the Pacific Ocean. The disjunct distribution of Australocarcinus is unexpected considering all trogoplacines are believed to practice direct development, lacking free-swimming larval stages. The new species is morphologically most similar to A. riparius Davie, 1988, from Queensland, Australia, but can be distinguished from its three congeners on the basis of the structures of its carapace, ambulatory legs and male first gonopod.

    Keywords: Chasmocarcinidae, freshwater, Indian Ocean, new species, Trogloplacinae, taxonomy

    Figure 1. Australocarcinus insperatus sp. n., holotype male (10.7 × 8.6 mm) (ZRC 2017.1072), Seychelles. A overall dorsal habitus B dorsal view of carapace (right side denuded) C right third maxilliped (denuded) D anterior thoracic sternum and pleon E posterior thoracic sternum and pleon F frontal view of cephalothorax G posterior margin of epistome. 

    Australocarcinus insperatus sp. n., holotype male (10.7 × 8.6 mm) (ZRC 2017.1072), Seychelles.
    Figure 1.  A overall dorsal habitus B dorsal view of carapace (right side denuded) C right third maxilliped (denuded) D anterior thoracic sternum and pleon E posterior thoracic sternum and pleon F frontal view of cephalothorax G posterior margin of epistome.
    Figure 2.  A outer surfaces of chelae B right first ambulatory leg showing setose posterior margin on propodus and dactylus C left fourth ambulatory leg D posterior thoracic sternum showing supplementary plate

    Figure 2. Australocarcinus insperatus sp. n. A–D holotype male (10.7 × 8.6 mm) (ZRC 2017.1072), Seychelles E–G paratype female (9.5 × 7.8 mm) (ZRC 2017.1073), Seychelles. A outer surfaces of chelae B right first ambulatory leg showing setose posterior margin on propodus and dactylus C left fourth ambulatory leg D posterior thoracic sternum showing supplementary plate E female overall dorsal habitus F female posterior thoracic sternum and pleon G female sterno-pleonal cavity showing vulvae.

    Family Chasmocarcinidae Serène, 1964
    Subfamily Trogloplacinae Guinot, 1986
    Genus Australocarcinus Davie, 1988
    Type species: Australocarcinus riparius Davie, 1988, by original designation.

    Australocarcinus insperatus sp. n.

    Material examined: Holotype: male (10.7 × 8.6 mm) (ZRC 2017.1072), in shallow stream, ca. 800 m from sea, about 2 km south-southeast of international airport, Mahé, Seychelles, coll. SR Daniels, May 2010. Paratypes: 1 male (8.5 × 7.2 mm), 1 female (9.5 × 7.8 mm) (ZRC 2017.1073), same data as holotype.

    Diagnosis: Carapace subquadrate, front weakly bilobed, with shallow median concavity (Fig. 1A, B); dorsal surface gently convex (Fig. 1F); dorsal surfaces and margins covered with short uneven tomentum (Fig. 1A, B); anterolateral margins arcuate, with four low teeth: first widest with gently sinuous margin, second lobiform, third wide, fourth (at junction of antero- and posterolateral margins) dentate, directed laterally, protruding beyond margin (Fig. 1B). Posterolateral margin converging towards gently convex posterior carapace margin (Fig. 1B). Epistome compressed, posterior margin with distinct triangular median lobe with median fissure, lateral margins gently sinuous (Fig. 1G). Eye peduncle completely filling orbit, relatively short, mobile; cornea distinct, pigmented (Fig. 1B, F). Third maxillipeds leaving gap when closed; merus quadrate, anteroexternal angle auriculiform; ischium quadrate, slightly longer than merus with very shallow median sulcus (Fig. 1C, D). Chelipeds subequal, relatively stouter in males (Figs 1A, 2E); cutting margins of both chelae with distinct teeth in both sexes, base of fingers with tuft of stiff setae; proximal part of dactylus of right chela with large, triangular tooth directed towards palm (Fig. 2A); ventral surface of cheliped merus with tubercles. Ambulatory legs moderately short; meri unarmed but setose to varying degrees; P2 carpus, propodus and dactylus with very long coarse setae which obscures margins (Figs 1A, 2B); P3–P5 propodus and dactylus setose but setae shorter than on P5 (Fig. 2C); P5 dactylus straight (Fig. 2C). Thoracic sternites 1, 2 fused, broadly triangular, short; separated from sternite 3 by sinuous groove; sternites 3, 4 fused, relatively broad (Fig. 1D). Male pleon with lateral margins of somite 6 and fused somites 3‒5 gently sinuous; telson slightly longer than broad (Fig. 1D, E). Sterno-pleonal cavity of male deep, press-button for pleonal holding small, short tubercle posterior to thoracic sternal suture 4/5 near edge of sterno-pleonal cavity. Male thoracic sternite 8 short, rectangular; supplementary plate narrow, wider along outer part (Figs 1E, 2D). G1 stout; basal part truncate; distal part cylindrical, with rounded tip, covered with short spinules (Fig. 3A–D). G2 prominently longer than G1, basal segment curved; distal segment slightly longer than basal segment, apex cup-like (Fig. 3E, F). Somites of female pleon with slightly convex lateral margins; telson wider than long (Fig. 2F). Sterno-pleonal cavity of female moderately deep, with large vulvae distinctly separated from each other, covering most of thoracic sternite 5, ovate, with low raised lip on outer margin, opening slit-like (Fig. 2G).

    Etymology: From the Latin “insperatus” for “unforeseen”, alluding to the unexpected discovery of a species of Australocarcinus in the western Indian Ocean.

     Peter K. L. Ng and Savel R. Daniels. 2018. A New Species of Trogloplacine Crab of the Genus Australocarcinus Davie, 1988 from A Freshwater Stream in Mahé, Seychelles (Crustacea, Brachyura, Chasmocarcinidae).  ZooKeys. 738; 27-35.   DOI:  10.3897/zookeys.738.23708

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    Hexanchus vitulus Springer & Waller, 1969

    in Daly-Engel, Baremore, Grubbs, et al., 2018. 

    The sixgill sharks of the genus Hexanchus (Hexanchiformes, Hexanchidae) are large, rarely encountered deep-sea sharks, thought to comprise just two species: the bluntnose sixgill Hexanchus griseus (Bonaterre, 1788) and the bigeye sixgill Hexanchus nakamurai (Teng, 1962). Their distribution is putatively worldwide in tropical and temperate waters, but many verified records for these species are lacking, and misidentification is common. Taxonomic uncertainty has long surrounded H. nakamurai in particular, with debate as to whether individuals from the Atlantic constitute a separate species. Using 1,310 base pairs of two mitochondrial genes, COI and ND2, we confirm that bigeye sixgill sharks from the Atlantic Ocean (Belize, Gulf of Mexico, and Bahamas) diverge from those in the Pacific and Indian Oceans (Japan, La Reunion, and Madagascar) with 7.037% sequence divergence. This difference is similar to the genetic distance between both Atlantic and Indo-Pacific bigeye sixgill sharks and the bluntnose sixgill shark (7.965% and 8.200%, respectively), and between the entire genus Hexanchus and its sister genus Heptranchias (8.308%). Such variation far exceeds previous measures of species-level genetic divergence in elasmobranchs, even among slowly-evolving deep-water taxa. Given the high degree of morphological similarity within Hexanchus, and the fact that cryptic diversity is common even among frequently observed shark species, we conclude that these results support the resurrection of the name Hexanchus vitulus Springer and Waller, 1969 for bigeye sixgill sharks in the northwest Atlantic Ocean. We propose the common name “Atlantic sixgill shark” for H. vitulus, and provide new locality records from Belize, as well as comments on its overall distribution.

    Keywords: Systematics, Mitochondrial DNA, Phylogenetics, Speciation, Elasmobranchs 

    An adult Atlantic sixgill shark swims in the waters off Belize.
    photo: Ivy Baremore/Maralliance

    Toby S. Daly-Engel, Ivy E. Baremore, R. Dean Grubbs, Simon J. B. Gulak, Rachel T. Graham and Michael P. Enzenauer. 2018. Resurrection of the Sixgill Shark Hexanchus vitulus Springer & Waller, 1969 (Hexanchiformes, Hexanchidae), with comments on its distribution in the northwest Atlantic Ocean. Marine Biodiversity.  DOI: 10.1007/s12526-018-0849-x

    New species of shark discovered through genetic testing via @physorg_com

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    Hyptiogaster arafura 
    Parslow & Jennings, 2018


    Hyptiogaster arafura sp. nov. is described from Arafura Swamp, Northern Territory, Australia, as the eleventh species of Hyptiogaster Kieffer (Hymenoptera: Gasteruptiidae). A revised diagnosis of Hyptiogaster is given based on the new species.

    Hyptiogaster arafura sp. nov. is described from Arafura Swamp, Northern Territory, Australia, as the eleventh species of Hyptiogaster. 

    Ben A. Parslow and John T. Jennings. 2018. A New Species of the Endemic Australian Genus Hyptiogaster Kieffer (Hymenoptera: Gasteruptiidae). Zootaxa. 4379(1); 145–150. DOI: 10.11646/zootaxa.4379.1.11

    The 11th species of an endemic Australian wasp genus via @physorg_com

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    Molossus fentoni
    Loureiro, Lim& Engstrom, 2018

    We describe a new species of mastiff bat in the genus Molossus (Molossidae), which was previously confused with the common and widely distributed M. molossus, from Guyana and Ecuador based on morphological and molecular differences. It is diagnosed by the following set of morphological characteristics: bicolored dorsal pelage, rounded anterior arch of the atlas, triangular occipital bone, and smaller body and skull size. In a molecular phylogenetic analysis of mitochondrial and nuclear DNA, maximum likelihood and parsimony trees recovered eight clades in the genus and a polyphyletic relationship for the M. molossus species complex. The new species was recovered in a well-supported clade that can be genetically distinguished from other species in the genus by its high level of sequence divergence based on the mitochondrial CO1 gene (8.0–10.1%) and on the nuclear gene beta fibrinogen (1.0–3.1%). It is broadly sympatric with M. molossus sensu stricto in northern South America, but morphologically distinct and genetically divergent.

     Keywords: Molossidae, New species, Phylogenetics, South America, Taxonomy

    Fig. 5. Holotype of Molossus fentoni sp. nov. (ROM 122583). Adult male with a medium brown dorsal pelage.

    Fig. 4. Dorsal, ventral, posterior, and lateral views of the skull of the holotype of Molossus fentoni sp. nov.

    Molossus fentoni sp. nov. 

    Diagnosis: A set of traits distinguishes Molossus fentoni from other Molossus. In M. fentoni the infra-orbital foramen is laterally directed; the basioccipital pits are rounded and of moderate depth; the occipital is triangular in posterior view; the upper incisors are thin and long with parallel tips and project forward in an oblique plane relative to the anterior face of the canines (Fig. 4); and the anterior arch of the atlas is rounded (Fig. 6).

    Distribution: Molossus fentoni is currently known from the administrative regions of Potaro-Siparuni and Upper Takutu-Upper Essequibo in Guyana and in Orellana province in Ecuador. Although, it has not been documented in the intervening 2000 km of lowland Amazonian forest, we anticipate that it will be found to have a broader distribution then initially represented in our collections. One individual of M. fentoni was collected in syntopy with M. coibensis, M. m. molossus, and M. rufus at ... east of Pompeya Sur, Orellana, Ecuador on 18 May, 2006.

    Etymology: This species is named in honour of M. Brock Fenton, Professor Emeritus, Western University, London, Ontario, and one of the world’s foremost researchers in bat ecology and behaviour. He was born in Guyana to Canadian parents and conducted fieldwork in the country in 1970.

    Taxonomic remarks: Husson (1962) designated the lectotype of M. molossus as the larger of the two bats described by Buffon and Daubenton (1763). Later, Husson (1962) restricted the type locality of M. molossus to Martinique, which previously had only been designated as the Americas in the first citation of this specimen (Buffon and Daubenton, 1759). Specimens of M. molossus from Martinique were morphologically analyzed in our study and have all the characteristics described above for M. molossus, and not for M. fentoni. In addition, the DNA sample of M. molossus from Martinique clustered with several other samples of M. molossus from the mainland in the phylogenetic trees (Fig. 2), such as Guyana, Suriname, and Brazil, corroborating its affiliation with M. molossus and the distinction from M. fentoni.

    Fig. 8. Schematic comparison of cranial features in Molossus.
    A and B – Posterior view; C and D – frontal view; E and F – Ventral view. Numbers represent characters described in the text. 1 – Lambdoidal crest and occipital complex; 2 – Sagittal crest; 3 – Mastoid process; 4 – Infra-orbital foramen; 5 – Upper incisors; 6 – Rostrum shape; 7 – Basioccipital pits.

     Livia O. Loureiro, Burton K. Lim and Mark D. Engstrom. 2018. A New Species of Mastiff Bat (Chiroptera, Molossidae, Molossus) from Guyana and Ecuador. Mammalian Biology. 90; 10-21.  DOI: 10.1016/j.mambio.2018.01.008 

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    Epicadus dimidiaster 
    Machado, Teixeira & Lise, 2018

    The Neotropical crab spider genera Tobias Simon, 1895 and Epicadus Simon, 1895 comprise species with remarkable somatic morphology and confounding taxonomic history. The results of our recent cladistic analysis corroborate and extend preceding taxonomic assumptions in showing that Tobias is a junior synonym of Epicadus. In the present paper the six species recently transferred from Tobias to Epicadus are redescribed. Two new species are described based on both males and females: Epicadus dimidiaster sp. nov. and Epicadus tigrinus sp. nov.; the male of Epicadus granulatus Banks, 1909 is described for the first time. The diagnosis of the genus is revised, an identification key is provided, and information on geographical distribution is updated. Epicadus now comprises eleven species.

    Keywords: Araneae, Dionycha, Neotropical region, Stephanopinae, taxonomy

    Miguel Machado, Renato Augusto Teixeira and Arno Antonio Lise. 2018. There and Back Again: More on the Taxonomy of the Crab Spiders Genus Epicadus (Thomisidae: Stephanopinae). Zootaxa. 4382(3);  501–530.  DOI:  10.11646/zootaxa.4382.3.4
    Thiago Da Silva Moreira and Miguel Machado. 2016. Taxonomic Revision of the Crab Spider Genus Epicadus Simon, 1895 (Arachnida: Araneae: Thomisidae) with Notes on Related Genera of Stephanopinae Simon, 1895. Zootaxa. 4147(3); 281–310.  DOI: 10.11646/zootaxa.4147.3.4

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    colonies of Pocillopora spp. from O‘ahu, Hawai‘i;
    (B–D) Pocillopora ligulata(F–I) P. meandrina and (K–M) P. eydouxi

    Johnston​, Forsman & Toonen, 2018.
     DOI:  10.7717/peerj.4355 

    Species within the scleractinian genus Pocillopora Lamarck 1816 exhibit extreme phenotypic plasticity, making identification based on morphology difficult. However, the mitochondrial open reading frame (mtORF) marker provides a useful genetic tool for identification of most species in this genus, with a notable exception of P. eydouxi and P. meandrina. Based on recent genomic work, we present a quick and simple, gel-based restriction fragment length polymorphism (RFLP) method for the identification of all six Pocillopora species occurring in Hawai‘i by amplifying either the mtORF region, a newly discovered histone region, or both, and then using the restriction enzymes targeting diagnostic sequences we unambiguously identify each species. Using this approach, we documented frequent misidentification of Pocillopora species based on colony morphology. We found that P. acuta colonies are frequently mistakenly identified as P. damicornis in Kāne‘ohe Bay, O‘ahu. We also found that P. meandrina likely has a northern range limit in the Northwest Hawaiian Islands, above which P. ligulata was regularly mistaken for P. meandrina.

    Figure 3: Images of Pocillopora ligulata colonies, (A)–(E); P. meandrina colonies, (F)–(J); and P. eydouxi colonies, (K)–(O) from O‘ahu, Hawai‘i. 

    Figure 1:Pocillopora species composition across the Hawaiian Islands for samples collected from colonies demonstrating P. meandrina morphology. The size of the pie chart is proportional to the number of individuals sampled per island. Pocillopora species are represented by different colors, specifically: P. meandrina, light yellow; P. eydouxi, dark yellow; P. ligulata, light blue; and P. verrucosa, dark blue.

    Here, we present an assay that allows rapid and unambiguous identification of all six species of Pocillopora present in Hawai‘i, which we hope will work anywhere these species are found. We present two cases where samples identified morphologically were misidentified to highlight the utility of this approach. Taxonomic confusion can impact a wide range of studies and the ability to rapidly and cost-effectively distinguish among species of Pocillopora will benefit future studies of population structure, ecology, biodiversity, evolution and conservation in this challenging genus.

    Erika C. Johnston​, Zac H. Forsman and Robert J. Toonen. 2018. A Simple Molecular Technique for Distinguishing Species reveals Frequent Misidentification of Hawaiian Corals in the Genus Pocillopora.  PeerJ. 6:e4355.  DOI:  10.7717/peerj.4355

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    Kaji, Anker, Wirkner & Palmer, 2018.

    • The evolutionary history of remarkable snapping claws in shrimp is reconstructed
    • Two novel claw-joint types—slip joints and torque-reversal joints—preceded snapping
    • The transition “slip joint → torque-reversal joint → snapping” occurred in two families
    • Subtle changes in joint form yielded dramatic changes in claw function (e.g., speed)

    How do stunning functional innovations evolve from unspecialized progenitors? This puzzle is particularly acute for ultrafast movements of appendages in arthropods as diverse as shrimps, stomatopods, insects, and spiders. For example, the spectacular snapping claws of alpheid shrimps close so fast (∼0.5 ms) that jetted water creates a cavitation bubble and an immensely powerful snap upon bubble collapse. Such extreme movements depend on (1) an energy-storage mechanism (e.g., some kind of spring) and (2) a latching mechanism to release stored energy quickly. Clearly, rapid claw closure must have evolved before the ability to snap, but its evolutionary origins are unknown. Unearthing the functional mechanics of transitional stages is therefore essential to understand how such radical novel abilities arise. We reconstructed the evolutionary history of shrimp claw form and function by sampling 114 species from 19 families, including two unrelated families within which snapping evolved independently (Alpheidae and Palaemonidae). Our comparative analyses, using micro-computed tomography (microCT) and confocal imaging, high-speed video, and kinematic experiments with select 3D-printed scale models, revealed a previously unrecognized “slip joint” in non-snapping shrimp claws. This slip joint facilitated the parallel evolution of a novel energy-storage and cocking mechanism—a torque-reversal joint—an apparent precondition for snapping. Remarkably, these key functional transitions between ancestral (simple pinching) and derived (snapping) claws were achieved by minute differences in joint structure. Therefore, subtle changes in form appear to have facilitated wholly novel functional change in a saltational manner.

    Keywords: Alpheidae, Palaemonidae, innovation, functional morphology, biomechanics, evolutionary morphology, evo-devo, comparative morphology, saltational evolution, torque-reversal joint

    Figure 1. MicroCT Images, Torque Moment Arms, and Schematic Illustrations of Three Shrimp Claw-Joint Types When Closed and Fully Open.
    (A) Pivot joint: anterior face∗ of right P1 in a basally branching caridean shrimp. (B) Simple slip joint (no torque reversal or power amplification): anterior face∗ of right P2 in an “intermediate” caridean shrimp. (C) Cocking slip joint (type 1 torque-reversal cocking, most likely power-amplified closing): anterior face∗ of right P1 in a feebly snapping alpheid shrimp. (A’–C’) Overlaid sagittal plane and surface rendering (via micro-computed tomography [microCT]) of claws of all three species showing torque moment-arms (+, –) when closed (upper) and fully opened (lower and background); negative torque (–) indicates that initial contraction of part of the closer muscle causes cocking. (A”–C”) Schematic representation of all three joint types showing loading orientations of opener and closer muscles. (A”) Pivot joint: purely rotational motion of dactyl. (B”) Slip joint: during opening, the dactylar base both rotates and translates (slips) across the propodal ridge (B). (C”) Cocking slip joint: during opening, the dactylar base both rotates and translates—including an abrupt sliding motion into the fully cocked position, where part of the closer muscle (gold) will generate reversed torque (–), and hence energy storage, because it inserts above the fulcrum (white dot).

     White dots show primary rotation axes (A–A”) or fulcrum points (B–B” and C–C”) for dactylar sliding and rotation. Black dots identify a reference point on the dactylar base. White arrows (A–C and A”–C”) show dactylar base trajectories during opening; closing would follow the same trajectories but in reverse. Red arrows (A’–C’) indicate dorsal-most closer-muscle contraction vectors (labeled V1 in Figure 4). Yellow arrows (A’–C’) represent torque moment arms about the fulcrum. Scale bars, 500 μm (A) and 300 μm (B and C). om, opener muscle; cm, closer muscle; (+), positive (counterclockwise) initial torque during claw closing; (–), negative (clockwise) torque during claw cocking generated by the gold-shaded muscle region in (C”). See also Figure SM1 in Methods S1 (joint-type scoring), Figures S1–S4 (microCT images of all claws), Movies S1A and S1B (actual dactyl motion), Movies S2A–S2E (3D model tests), and Table S1 (joint types of all species). ∗See Supplemental Results (Methods S1) for an explanation of claw-face viewing perspectives.

     Tomonari Kaji, Arthur Anker, Christian S. Wirkner and A. Richard Palmer. 2018. Parallel Saltational Evolution of Ultrafast Movements in Snapping Shrimp Claws. Current Biology.  28(1); 106-113.  DOI: 10.1016/j.cub.2017.11.044 

    An adaptation 150 million years in the making via @physorg_com

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    Aplocheilus andamanicus  (Köhler, 1906) 

    in Katwate, Kumkar, Britz, Raghavan & Dahanukar, 2018. 


    In his work on the fishes of the Andaman Islands, Francis Day (1870) collected large-sized specimens of Aplocheilus from the south Andamans. Despite differences in the size and dorsal-fin ray counts, Day refrained from recognising the Andaman Aplocheilus as a distinct species and considered it as Aplocheilus panchax, a species distributed in the Ganges delta and across the eastern coast of mainland India. However, Day mentioned the differences in fin-ray counts between these two populations. Subsequently Köhler (1906) described the Andaman population as Haplochilus andamanicus (now in Aplocheilus), referring to the diagnostic characters initially discovered by Day. This species failed to receive recognition from taxonomists, because of the uncertainty regarding the validity of the species and its questionable synonymy with A. panchax. In this study, based on morphological and molecular evidence, we demonstrate that A. andamanicus is indeed a distinct and valid species, which can easily be diagnosed from the widespread A. panchax. While resolving the identity of A. andamanicus, we also demonstrate that the congeners from southeast Asia form a genetically distinct group for which the name Odontopsis armata is available.

    Keywords: Pisces, Aplocheilus panchax, freshwater fish, taxonomy, South Asia


    Unmesh Katwate, Pradeep Kumkar, Ralf Britz, Rajeev Raghavan and Neelesh Dahanukar. 2018. The Identity of Aplocheilus andamanicus (Köhler, 1906) (Teleostei: Cyprinodontiformes), An Endemic Killifish from the Andaman Islands, with Notes on Odontopsis armata van HasseltZootaxa. 4382(1); 159–174.  DOI:  10.11646/zootaxa.4382.1.6


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    Acantholimon ibrahimii  Akaydın

    in Akaydın, 2018.


    A new species, Acantholimon ibrahimii Akaydın, is described, illustrated and discussed in comparison with its close relative A. davisii. The new species is distinguished from the latter species mainly by the generative organs (namely the inflorescence types and petals colour), habitat type and ecological behaviour. Data are also reported on the conservation status of A. ibrahimii, which is suggested to be labelled as EN according to the IUCN categories. Furthermore, a revised key to the Turkish Acantholimon species of A. sect. Staticopsis with spike laxly distichous and scape much longer than leaves is presented.

    Keywords: Acantholimon, A. sect. Staticopsis, conservation, endemism, Staticoideae, taxonomy, Eudicots

    Galip Akaydın. 2018. Acantholimon ibrahimii (Plumbaginaceae), A New Species of A. section Staticopsis from the Mediterranean Part of Turkey. Phytotaxa. 340(1); 48–54. DOI: 10.11646/phytotaxa.340.1.2

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    Thismia neptunis Beccari

    in Sochor, Egertová, Hroneš & Dančák, 2018. 

    Thismia neptunis, as many of its congeners, is a poorly understood species that has only been known from the type collection and its limited original description. In January 2017 it was rediscovered in the type area in the Gunung Matang massif, western Sarawak, Borneo, Malaysia. The paper provides the amended description and drawings of the species, very first available photographs and short notes on taxonomy and historical context of Beccari’s work on Thismia

    Key words: Brunonithismia, Burmanniaceae, fairy lanterns, Kubah, Monte Mattán, Sarawak

    FIGURE 3. Thismia neptunis: flowering plants (A, B), bud (C), detail of flower (D), section of floral tube and outer view of connective tube (E), detail of inner perianth lobe (F), stigma (G), lateral appendage (H).

    Taxonomic treatment 
    Thismia neptunis Beccari (1878: 251)

     Type:—MALAYSIA. Ragiato di Sarawak, Mattang. April 1866. O. Beccari p.b. 1508 (holotype FI-B 013453!)

    Habitat and ecology:—The only known locality is in primary lowland mixed dipterocarp forest on a river alluvium. Thismia species are generally accompanied by other mycoheterotrophic plants; in this case it was Sciaphila cf. alba Tsukaya & Suetsugu (2015: 284). Albeit pollination ecology was not studied, ca. seven flies of family Sciaridae (Diptera) and one individual of family Braconidae (Hymenoptera) were observed being stuck on inner perianth lobes of the two flowers (Fig. 3A, D, E, F). Although the braconid was probably only a coincidental victim, the flies may represent potential pollinators, as several dipteran taxa have been reported as visitors and probable pollinators of fairy lanterns (Li & Bi 2013, Mar & Saunders 2015). Nevertheless, why had they been attracted to and finally trapped on the perianth lobes surface can only be speculated. Tepals are apparently hydrophilic (possibly as a mean of maintaining turgor in the long thin appendages) as indicated by a number of rain drops persisting on them long after the rain. But they do not appear to be sticky and no other particles tended to be trapped on them either in the field or during our manipulation. Therefore, the insects seem to have been attracted by smell (or other signals) of the flowers and accidentally drowned on the wet surface of perianth lobes.

    Distribution:—The species is known from a restricted area in western Sarawak, Borneo, Malaysia. Beccari (1878) described the locality simply as “Monte Mattán” or “Mattang”, which is an area now generally known as Matang massif which Kubah National Park is part of it. The present locality is placed at the park’s western border and may be identical or close to that of Beccari.

    Taxonomic affinities:—Having free perianth lobes of unequal length and shape, T. neptunis belongs to section Thismia (Euthismia Schlechter, 1921: 34), subsection Brunonithismia Jonker (1938: 242). This group comprises nine species (Kumar et al. 2017, Suetsugu et al. 2018) of very diverse morphology as for symmetry of perianth, modification of perianth lobes and structure of connectives. Half of the species are, nevertheless, only poorly documented. Thismia neptunis is unique among other fairy lanterns in the very complex three-segmental structure of inner perianth lobes that are terminated by long filiform appendage pointing vertically upwards. This striking morphology led Schlechter to creation of monotypic section Sarawakia Schlechter (1921: 35) within his system of Thismia (Schlechter 1921). However, his approach has not been generally accepted (Jonker 1938, Kumar et al. 2017). 

    Beccari was also well aware of morphological uniqueness of T. neptunis. In the protologue (Beccari 1878), he stated that T. neptunis seems to have connectives similar to T. brunonis Griffith (1844: 221). However, T. brunonis have apical part of the connective covered by numerous short teeth (Griffith 1845) while T. neptunis have only three rather long appendages. Nevertheless, Beccari himself was not absolutely sure about the character of connectives as he studied only two pressed and dried plants. In having whitish perianth tube with 12 orange streaks T. neptunis superficially resembles T. javanica Smith (1910: 32) and T. arachnites Ridley (1905: 197). Both of them, nevertheless, differ in having short rounded outer perianth lobes and simpler spreading inner perianth lobes, and the latter species also in having “numerous short teeth” at the apical end of connectives. Connectives of T. javanica, although similar at a first glance, differ from those of T. neptunis in colour (white vs. orange, respectively) and three short teeth at the apex, each bearing 1–2 long hairs of similar length (vs. three unequal filiform appendages in T. neptunis). Thismia neptunis is so far the only known member of subsection Brunonithismia occurring in Borneo.

    Michal Sochor, Zuzana Egertová, Michal Hroneš and Martin Dančák. 2018. Rediscovery of Thismia neptunis (Thismiaceae) After 151 Years. Phytotaxa. 340(1); 71–78.  DOI: 10.11646/phytotaxa.340.1.5

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    Uperodon rohani
    Garg, Senevirathne, Wijayathilaka, Phuge, Deuti, Manamendra-Arachchi, Meegaskumbura & Biju, 2018

       DOI:  10.11646/zootaxa.4384.1.1 


    Based on a recent molecular phylogenetic study, the South Asian microhylid genus Uperodon (subfamily Microhylinae) currently comprises of 12 valid species that are largely restricted to India and Sri Lanka. Considering the revised generic-level status of its various members, here we review the taxonomy of all known species in this genus and clarify their nomenclatural status and geographical distribution, by integrating evidence from genetics, adult and tadpole morphology, breeding ecology, and bioacoustics. Our molecular analyses of a mitochondrial 16S rRNA gene fragment combined with external and internal morphological studies also revealed a distinct new species in the genus. This species, formally described as Uperodon rohani sp. nov., is endemic to Sri Lanka and widely distributed at lower elevations in the island. For nomenclatural stability of various previously known members, the following actions are also undertaken: (1) redescription of the poorly-defined species Ramanella anamalaiensis Rao (= Uperodon anamalaiensis) and Hylaedactylus montanus Jerdon (= Uperodon montanus); (2) neotype designation for Ramanella anamalaiensis Rao (= Uperodon anamalaiensis), Ramanella minor Rao, Ramanella mormorata Rao (= Uperodon mormorata), and Ramanella triangularis rufeventris Rao; (3) lectotype designation for Callula variegata Stoliczka (= Uperodon variegatus); and (4) synonymization of Ramanella minor Rao with Uperodon anamalaiensis.

    Keywords: Amphibia, Amphibians, bioacoustics, endemism, mitochondrial DNA, natural history, neotype, lectotype, new species, tadpoles, Western Ghats-Sri Lanka biodiversity hotspot

    Uperodon palmatus (Parker, 1934)

    Sonali Garg, Gayani Senevirathne, Nayana Wijayathilaka, Samadhan Phuge, Kaushik Deuti, Kelum Manamendra-Arachchi, Madhava Meegaskumbura and SD Biju. 2018. An Integrative Taxonomic Review of the South Asian Microhylid Genus UperodonZootaxa.  4384(1); 1–88.  DOI:  10.11646/zootaxa.4384.1.1

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    Hyobanche hanekomii  A. Wolfe
      H. atropurpurea Bolus
      H. sanguinea L. 

    in Wolfe, 2018. 


    The new species Hyobanche hanekomii is described and illustrated. It is somewhat intermediate in appearance between H. sanguinea and H. atropurpurea, but can be distinguished from both in several morphological characters that are presented. The new species occurs in the Cape Fold Belt Mountains of the northwest part of the Western Cape.

     Key words: Cape Floristic Region, holoparasite, parasitic plant

    FIGURE 3. Comparison of Hyobanche hanekomii (A), H. atropurpurea (B), and H. sanguinea (C).
    Upper panel represents side and front views of corollas to scale (bar = 10 mm).
    Lower panel shows inflorescences (not to scale). Illustrations and photographs by A. Wolfe; photos are from living plants in prime blooming condition. Vouchers: (A) upper panel: Hanekom 2887 (1997, NBG); lower panel: Wolfe 1009 (2001, OS); (B) Wolfe 1227 (2006, OS); (C) Wolfe 1387 (2013, OS). 

    Hyobanche hanekomiiA. Wolfe spec. nov.
    . Corolla deep magenta to magenta-red, inflated above the tube and semi-galeate, and 1.5–2.0 times the length of the calyx. 

    TYPE:—South Africa. Western Cape: ..., 400 m, 26 September 1997, W.J. Hanekom 2887 (Holotype: NBG 759260!) 

    Distribution. Rocky soils in Cape Fold Belt Mountains of northwestern region of the Western Cape, from Citrusdal area to Giftberg (Fig. 2). 

    Etymology. The specific epithet is in honour of Mr. Willem Johannes Hanekom (b. 1931). Mr. Hanekom is a keen observer of the flora of the Western Cape, and introduced the author to this new species in 2001. He had made a collection in 1997 (W. J. Hanekom 2887), which included the following note: “Hyobanche sanguinea L. but with influence of H. atropurpurea H. Bol.” 

    Andrea D. Wolfe. 2018. Hyobanche hanekomii (Orobanchaceae), A New Species from the Western Cape of South Africa. Phytotaxa. 340(1); 93–97.  DOI: 10.11646/phytotaxa.340.1.5

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    Siamophryne troglodytes
    Suwannapoom, Sumontha, Tunprasert, Ruangsuwan, Pawangkhanant, Korost & Poyarkov, 2018

    Tenasserim Cave Frog  • อึ่งถ้ำตะนาวศรี   |   DOI: 10.7717/peerj.4422 


    We report on a discovery ofSiamophryne troglodytes Gen. et sp. nov., a new troglophilous genus and speciesof microhylid frog from a limestone cave in the tropical forests of western Thailand. To assess its phylogenetic relationships we studied the 12S rRNA–16S rRNA mtDNA fragment with final alignment comprising up to 2,591 bp for 56 microhylid species. Morphological characterization of the new genus is based on examination of external morphology and analysis of osteological characteristics using microCT-scanning. Phylogenetic analyses place the new genus into the mainly Australasian subfamily Asterophryinae as a sister taxon to the genus Gastrophrynoides, the only member of the subfamily known from Sundaland. The new genus markedly differs from all other Asterophryinae members by a number of diagnostic morphological characters and demonstrates significant mtDNA sequence divergence. We provide a preliminary description of a tadpole of the new genus. Thus, it represents the only asterophryine taxon with documented free-living larval stage and troglophilous life style. Our work demonstrates that S. troglodytes Gen. et sp. nov. represents an old lineage of the initial radiation of Asterophryinae which took place in the mainland Southeast Asia. Our results strongly support the “out of Indo-Eurasia” biogeographic scenario for this group of frogs. To date, the new frog is only known from a single limestone cave system in Sai Yok District of Kanchanaburi Province of Thailand; its habitat is affected by illegal bat guano mining and other human activities. As such, S. troglodytes Gen. et sp. nov. is likely to be at high risk of habitat loss. Considering high ecological specialization and a small known range of the new taxon, we propose a IUCN Red List status of endangered for it.

    Keywords: Kanchanaburi Province, Siamophryne troglodytes Gen. et sp. nov., Tadpole, Troglophilous life style, Tenasserim, Sundaland, mtDNA, Biogeography, microCT-scanning

    Based upon the results of phylogenetic analyses of 12S rRNA–16S rRNA mtDNA fragment sequences, the Microhylidae frog from Kanchanaburi Province represents a previously unknown highly divergent mtDNA lineage, clearly distinct from all other members of Microhylidae for which comparable genetic data were available. This lineage falls into the Australasian subfamily Asterophryinae and with high values of node support is reconstructed as a sister group to the genus Gastrophrynoides that inhabits Borneo and the Peninsular Malaysia. Subsequent analyses of osteology and external morphology (see below) clearly indicate that the recently discovered population of Microhylidae Gen. sp. from Kanchanaburi Province represents a new previously undescribed genus and species which we describe herein as:

    Amphibia Linnaeus, 1758
    Anura Fischer von Waldheim, 1813

    Microhylidae Günther, 1858
    Asterophryinae Günther, 1858

    Figure 5: Male paratype of Siamophryne troglodytes Gen. et sp. nov. (ZMMU A-5818) in life in dorsolateral aspect. Photo by N. A. Poyarkov.

    Siamophryne Gen. nov.

    Diagnosis: A medium-sized (19 mm < SVL < 30 mm) member of the Australasian subfamily Asterophryinae (family Microhylidae), with the following combination of morphological attributes: (1) both maxillae and dentaries eleutherognathine, no maxillary teeth; (2) vertebral column procoelous with eight presacral vertebrae (PSV) lacking neural crests; (3) no sagittal crest on cranium; (4) frontoparietals conjoined, connected by long suture; (5) nasals wide, calcified, but not contacting each other medially; (6) vomeropalatines small, not expanded, vomerine spikes absent; (7) cultriform process of parasphenoid comparatively narrow; (8) clavicles present as slender tiny bones, lying on the procoracoid cartilage not reaching scapula or the midline; (9) omosternum absent; (10) sternum large, anterior portion consists of calcified cartilage, xiphisternum cartilaginous; (11) weak dorsal crest present on urostyle, absent on ilium; (12) terminal phalanges large T-shaped; (13) all fingers and toe discs with terminal grooves; (14) subarticular tubercles weak, discernible only at digit basis; (15) toe webbing absent; (16) tympanum distinct; (17) two transverse smooth palatal folds; (18) pupil round; (19) snout rounded, equal to EL; (20) development with a larval stage, tadpole with peculiar dorso-ventrally compressed morphology.

    Type species. Siamophryne troglodytes sp. nov.
    Other included species. None are known at present.

    Distribution: To date, S. troglodytes sp. nov. is only known from a small cave system in a karst region of Sai Yok District, Kanchanaburi Province, northern Tenasserim Region, western Thailand (see below the description of the species) (see Fig. 1).

    Etymology: The generic nomen Siamophryne is derived from “Siam”—the old name of present-day Thailand; referring to the range of the new genus, which to date is only known from western Thailand; and the Greek noun “phryne” (φρÚνη; feminine gender), meaning “toad” in English; this root is often used in the generic names in Asterophryinae microhylid frogs. Gender of the new genus is feminine.

    Figure 10: Breeding habitat of Siamophryne troglodytes Gen. et sp. nov. at the type locality—Sai Yok District, Kanchanaburi Province, northern Tenasserim Region, western Thailand.  (A) Entrance to the limestone cave where the frogs were recorded; (B) female in situ sitting on the limestone wall of the cave; (C) male in situ sitting in a water-filled crevice; (D) female in situ on the wall of the cave (photos by M. Sumontha);  


    Figure 10: Breeding habitat of Siamophryne troglodytes Gen. et sp. nov. at the type locality—Sai Yok District, Kanchanaburi Province, northern Tenasserim Region, western Thailand.
     (A) Entrance to the limestone cave where the frogs were recorded; (B) female in situ sitting on the limestone wall of the cave; (C) male in situ sitting in a water-filled crevice; (D) female in situ on the wall of the cave (photos by M. Sumontha); (E, F) tadpole in situ in a water-filled crevice (photos by T. Ruangsuwan).

    Figure 8: Tadpole of  Siamophryne troglodytes Gen. et sp. nov. in life (AUP-00509; Gosner stage 36).
     (A) In dorsal and (B) in ventral aspects. Scale bar equals to 5 mm. Photos by N. A. Poyarkov. Tadpole of Siamophryne troglodytes Gen. et sp. nov. in preservative (AUP-00509; Gosner stage 36). 

    Figure 9: Tadpole of Siamophryne troglodytes Gen. et sp. nov. in preservative (AUP-00509; Gosner stage 36). (A) In lateral, (B) in dorsal, and (C) in ventral views. Scale bar equals to 5 mm. Photos by T. Ruangsuwan.

    Figure 10: Breeding habitat of Siamophryne troglodytes Gen. et sp. nov. at the type locality—Sai Yok District, Kanchanaburi Province, northern Tenasserim Region, western Thailand.
     (C) male in situ sitting in a water-filled crevice; (D) female in situ on the wall of the cave (photos by M. Sumontha); (E, F) tadpole in situ in a water-filled crevice (photos by T. Ruangsuwan).

    Siamophryne troglodytes sp. nov.

    Etymology: The specific name “troglodytes” is a Latin adjective in the nominative singular meaning “cave-dweller”, derived from the Greek “τρωγλoδύτης”, with “trogle” meaning “holemouse-hole” and “dyein” meaning “go indive in”; referring to the troglophilous biology of the new species, which was recorded only in a limestone karst cave system.

    Suggested common names: We recommend the following common names for the new species: “Tenasserim Cave Frog” (English); “อึ่งถ้ำตะนาวศรี - Eung Tham Tenasserim” (Thai).

    Natural history notes:  
    Siamophryne troglodytes Gen. et sp. nov. has a troglophilous life style and to date is only known from a small limestone cave system in western Thailand. All specimens were collected within a narrow area inside a limestone cave located on elevation 440 m a.s.l. in a polydominant tropical forest in Sai Yok District, Kanchanaburi Province, western Thailand (Fig. 10A). The cave was examined twice on the 1st of August and the 27th of October, 2016. In both cases, adult specimens of S. troglodytes Gen. et sp. nov. were only recorded inside the cave, at a distance of more than 25 m from the entrance, sitting on walls of the cave (Figs. 10B and 10D) or hiding inside small caverns in limestone (Fig. 10C) or under flat stones. Despite the thorough search, no animals were recorded near the cave entrance or in the forest close to the cave. Animals were active from 23:00 to 24:00, when the air temperature inside the cave was 28 °C in August and 26 °C in October, in both cases with 100% humidity. No calling activity was recorded during both surveys. Diet and enemies of the new frog are unknown.

    Three tadpoles (one of which was collected) were observed during the survey on the 1st of August, 2016, in a small water-filled cavity in the limestone on the floor of the cave, ca. 10 m from the cave entrance (Figs. 10E and 10F). The cavity was filled with water, the average depth was 4–5 cm; mosquito larvae (Chironomidae) were also observed in the same water body. Four other tadpoles (not collected) were discovered in another similar water-filled cavity inside the cave (30 m from the cave entrance).

    The cave system where S. troglodytes Gen. et sp. nov. was discovered is inhabited by several species of bats which produce significant amount of guano that accumulates on the cave floor. According to a local guide, the locals mine this guano and that affects the ecosystem of the cave.

    Distribution: As for the genus. At present, S. troglodytes Gen. et sp. nov. is known from a single limestone karst cave in Sai Yok District of Kanchanaburi Province in western Thailand. To date, numerous surveys in the nearby karst massifs have not yielded discoveries of additional populations of the new species. However, further fieldwork in Kanchanaburi Province of Thailand and the adjacent parts of Tanintharyi Division of Myanmar are required.

    Siamophryne troglodytes, a new genus and species of microhylid frogs from western Thailand, belongs to the subfamily Asterophryinae, which is most diverse in Australasia. Siamophryne and its sister genus Gastrophrynoides are the only two asterophryine lineages found in the areas derived from the Eurasian landmass. Our work demonstrates that S. troglodytes represents an old lineage of the initial radiation of Asterophryinae which took place in the mainland Southeast Asia. Our results strongly support the “out of Indo-Eurasia” biogeographic scenario for this group of frogs. To date, the new frog is the only known asterophryine with a free-living tadpole and troglophilous life style. Further studies might reveal new members of Asterophryinae in the mainland Southeast Asia.

     Chatmongkon Suwannapoom, Montri Sumontha, Jitthep Tunprasert, Thiti Ruangsuwan, Parinya Pawangkhanant, Dmitriy V. Korost and Nikolay A. Poyarkov. 2018. A Striking New Genus and Species of Cave-dwelling Frog (Amphibia: Anura: Microhylidae: Asterophryinae) from Thailand.  PeerJ. 6:e4422.  DOI: 10.7717/peerj.4422