[Paleontology • 2015] Probrachylophosaurus bergei • A New Brachylophosaurin Hadrosaur (Dinosauria: Ornithischia) with an Intermediate Nasal Crest from the Campanian Judith River Formation of Northcentral Montana

Probrachylophosaurus bergei 

Fowler & Horner, 2015

  Illustration: John Conway  JohnConway.co || DOI: 10.1371/journal.pone.0141304

Abstract

Background

Brachylophosaurini is a clade of hadrosaurine dinosaurs currently known from the Campanian (Late Cretaceous) of North America. Its members include: Acristavus gagslarsoni, which lacks a nasal crest; Brachylophosaurus canadensis, which possesses a flat paddle-shaped nasal crest projecting posteriorly over the dorsal skull roof; and Maiasaura peeblesorum, which possesses a dorsally-projecting nasofrontal crest. Acristavus, from the lower Two Medicine Formation of Montana (~81–80 Ma), is hypothesized to be the ancestral member of the clade. Brachylophosaurus specimens are from the middle Oldman Formation of Alberta and equivalent beds in the Judith River Formation of Montana; the upper Oldman Formation is dated 77.8 Ma.

Methodology/Principal Findings

A new brachylophosaurin hadrosaur, Probrachylophosaurus bergei (gen. et sp. nov.) is described and phylogenetically analyzed based on the skull and postcranium of a large individual from the Judith River Formation of northcentral Montana (79.8–79.5 Ma); the horizon is equivalent to the lower Oldman Formation of Alberta. Cranial morphology of Probrachylophosaurus, most notably the nasal crest, is intermediate between Acristavus and Brachylophosaurus. In Brachylophosaurus, the nasal crest lengthens and flattens ontogenetically, covering the supratemporal fenestrae in large adults. The smaller nasal crest of Probrachylophosaurus is strongly triangular in cross section and only minimally overhangs the supratemporal fenestrae, similar to an ontogenetically earlier stage of Brachylophosaurus. Sutural fusion and tibial osteohistology reveal that the holotype of Probrachylophosaurus was relatively more mature than a similarly large Brachylophosaurus specimen; thus, Probrachylophosaurus is not simply an immature Brachylophosaurus.

Conclusions/Significance

The small triangular posteriorly oriented nasal crest of Probrachylophosaurus is proposed to represent a transitional nasal morphology between that of a non-crested ancestor such as Acristavus and the large flat posteriorly oriented nasal crest of adult Brachylophosaurus. Because Probrachylophosaurus is stratigraphically and morphologically intermediate between these taxa, Probrachylophosaurus is hypothesized to be an intermediate member of the Acristavus-Brachylophosaurus evolutionary lineage.

Systematic Paleontology

Dinosauria Owen, 1842

Ornithischia Seeley, 1888

Ornithopoda Marsh, 1881

Hadrosauridae Cope, 1869

Hadrosaurinae Cope, 1869

Brachylophosaurini Gates et al., 2011

Definition: Modified from Gates et al. [3]: Hadrosaurine ornithopods more closely related to Brachylophosaurus, Probrachylophosaurus, Maiasaura, or Acristavus than to Gryposaurus or Saurolophus.

Referred material: UCMP 130139, a partial skull and skeleton originally described as the holotype of Brachylophosaurus goodwini [9], and later assigned to Brachylophosaurus canadensis [1, 23]. Due to the lack of a preserved nasal, and the presence of deep frontal depressions, the specimen cannot be confidently assigned to any current genus of Brachylophosaurini.

Horizon and locality: UCMP 130139 was collected from the Judith River Formation of Kennedy Coulee, Hill County, northcentral Montana, in beds equivalent to the lower Oldman Formation, with a published height of approximately 15 m above the Marker A Coal of the Taber Coal Zone of the Foremost Formation [9]. However, a remeasured section shows that the site was actually only a few meters above the Marker A Coal, and lies within the Herronton Sandstone Zone (Mark Goodwin and David Evans personal communication, 2014).

Brachylophosaurus canadensis Sternberg, 1953

Holotype: CMN 8893

Referred Material: FMNH PR 862 (partial skull); MOR 720 (braincase); MOR 794 (nearly complete articulated skeleton); MOR 940 (braincase); MOR 1071 (monodominant bonebed); TMP 90.104.01 (complete skull and articulated partial skeleton).

Horizons and localities: CMN 8893, FMNH PR 862, and TMP 90.104.01 were collected from the Oldman Formation of southeastern Alberta. CMN 8893 and FMNH PR 862 were collected in Dinosaur Provincial Park; TMP 90.104.01 was collected near Onefour and the Milk River. Of these Albertan specimens, the exact stratigraphic position is known only for CMN 8893: the Comrey Sandstone Zone (Unit 2) of the Oldman Formation. MOR 720 was collected from the upper Judith River Formation in badlands surrounding the Missouri River north of Winifred, Fergus County, central Montana. MOR 794, MOR 940, and MOR 1071 were collected from the Judith River Formation of Malta, northern Montana, in beds equivalent to the Comrey Sandstone Zone of the Oldman Formation.

Probrachylophosaurus gen. nov.

urn:lsid:zoobank.org:act:7B7C87AC-2EFE-4587-9A24-A5D48C908941

Etymology: Pro- (Latin) before, -brachylophosaurus (Greek) short-crested lizard, in reference to the new taxon’s stratigraphic position below that of Brachylophosaurus canadensis.

Probrachylophosaurus bergei sp. nov.

urn:lsid:zoobank.org:act:49D503CB-7FA6-4D66-8FC0-0B4E2A3EE106

Etymology: Species name bergei in memory of Sam Berge, co-owner of the land where the specimen was discovered, and friend and relative of many members of the Rudyard, Montana community, who have supported paleontologic research for decades. Pronunciation: berg-ee-i

Conclusions

In the early years of dinosaur paleontology, specimens were collected as isolated points, each so morphologically unique that their evolutionary relationships were difficult to determine. As more fossil specimens are collected, the gaps in morphology that previously separated species are being filled. With larger sample sizes resulting in more continuous series of fossils with good stratigraphic resolution, variations in morphology can be analyzed in a more complete context, and attributed to ontogeny, evolution, taphonomic alteration, biogeography, or individual variation. A better understanding of stratigraphy and advancements in radiometric dating enable precise temporal correlation of geographically separated localities. Taxa can then be placed in temporal sequence, allowing tests of evolutionary hypotheses.

A precise stratigraphic framework is critical for determining whether morphological variations of adult specimens are due to evolution or are variations within a roughly contemporaneous population. If closely related taxa do not overlap stratigraphically, the pattern is more parsimonious with anagenesis than cladogenesis. Recent research has greatly increased the sample size and stratigraphic resolution of specimens from the Judith River and Hell Creek Formations of Montana and their Canadian equivalents, revealing several potential anagenetic lineages in Campanian and Maastrichtian ornithischians.

Because Probrachylophosaurus bergei is stratigraphically older than all Brachylophosaurus canadensis specimens, it is hypothesized to represent a basal brachylophosaur morphology, early in the evolution of this lineage from a non-crested ancestor. Thus, the small crest of Probrachylophosaurus would represent a transitional nasal morphology between a non-crested ancestor such as Acristavus and the larger crests of adult Brachylophosaurus. The fourth member of Brachylophosaurini, Maiasaura, would represent a cladogenic event, diverging from the lineage that led to Brachylophosaurus at a currently unknown point.

Elizabeth A. Freedman Fowler and John R. Horner. 2015. A New Brachylophosaurin Hadrosaur (Dinosauria: Ornithischia) with an Intermediate Nasal Crest from the Campanian Judith River Formation of Northcentral Montana. PLoS ONE. DOI: 10.1371/journal.pone.0141304  @PLOSPaleo @theJohnConway

MSU News - MSU team finds new dinosaur species, reveals evolutionary link: http://www.montana.edu/news/15858/msu-team-finds-new-dinosaur-species-reveals-evolutionary-link

New 'short-crested lizard' found in Montana http://phy.so/366472776 via @physorg_com

[Herpetology / Behaviour • 2015] Avian Deception Using An Elaborate Caudal Lure in Pseudocerastes urarachnoides (Serpentes: Viperidae)

ABSTRACT

  Pseudocerastes urarachnoidesis a fascinating viper and as yet has been reported only in western Iran. An elaborated arachnid-like caudal structure is a unique feature of this viper, hence gives it the common name “Iranian spider-tailed viper”. During tail wagging, the structure is reminiscent of a moving spider. Tail movements are used for two different purposes in snakes: defense via tail vibration and hunting via both caudal luring and caudal distraction. Caudal luring in snakes is the wriggling or wagging of the posterior part of tail, in the presence of a potential prey, with conspicuous color pattern while the rest of body is cryptically colored. Previous studies have speculated on the role of caudal structure of P. urarachnoides in hunting. Our 2.5-year study has revealed that development of the structure of the caudal lure is commenced after birth and is linearly correlated to snout-vent length. The caudal lure attracts some species of birds. Caudal luring behavior is carried out both in the presence and absence of birds. The findings are reported for the first time and confirmed by direct observation of undisturbed individuals in the field.

  Keywords: caudal luring, Iranian spider-tailed viper, prey attraction, Pseudocerastes urarachnoides.

http://figshare.com/articles/Avian_deception_using_an_elaborate_caudal_lure_in_Pseudocerastes_urarachnoides_Serpentes_Viperidae_/1454446

 Behzad Fathinia, Nasrullah Rastegar-Pouyani, Eskandar Rastegar Pouyani, Fatemeh Todehdehghan and Fathollah Amiri. 2015. Avian Deception Using An Elaborate Caudal Lure in Pseudocerastes urarachnoides (Serpentes: Viperidae).
AMPHIBIA-REPTILIA. 36(3); 223 – 231. DOI:  10.1163/15685381-00002997

ResearchGate.net/publication/280348452_Avian_deception_using_an_elaborate_caudal_lure_in_Pseudocerastes_urarachnoides_(Serpentes_Viperidae)

[Herpetology • 2015] Revision of the Western Australian Pebble-mimic Dragon Species-Group (Tympanocryptis cephalus: Reptilia: Agamidae)

FIGURE 5. Photographs in life of Tympanocryptis cephalus species-group members (left to right, top to bottom):
 a) T. cephalus (photo—G. Harold), b) T. gigas (M. Peterson), c) T. diabolicus sp. nov. (M. Peterson), d) T. fortescuensis sp. nov. (G. Harold), e) T. pseudopsephos sp. nov. —plain form (G. Harold), f) T. pseudopsephos sp. nov.—with charcoal wash on dorsum (G. Harold).

DOI:  10.11646/zootaxa.4039.1.3

Abstract

Recent work on species complexes of the pebble-mimic dragons of the Australian genus Tympanocryptis has greatly clarified evolutionary relationships among taxa and also indicated that species diversity has been severely underestimated. Here we provide a morphological and molecular appraisal of variation in the T. cephalus species-group and find evidence for recognizing five species-level lineages from Western Australia. Four species-level lineages are strongly supported with a combined mitochondrial and nuclear DNA Bayesian analysis (a fifth population from the Gascoyne region lacked tissue samples). Morphologically, we found subtle, yet consistent, differences among the populations in scalation, color and pattern. True T. cephalus Günther is restricted to the coastal Pilbara region and characterized by five dark blotches on the dorsum, keeled ventrals, and other characters. Two other lineages within the Pilbara, from the Hamersley range and Fortescue/northern Pilbara region, differed from T. cephalus senso stricto by possessing a more elongate body and a plain dorsum. Furthermore, the Hamersley lineage differed from the Fortescue lineage by possessing slightly more reddish coloration and feeble keeling on the snout. Although there are few specimens and no tissue samples available for the Gascoyne population, these individuals are larger, have rugose scales on the snout, and possess scattered enlarged tubercles with three large blotches on the dorsum. The name T. cephalus gigas Mitchell is available for this population. The most widespread lineage, and the one best represented in collections and in field guides, occurs throughout central Western Australia. These Goldfield populations are characterized by a protruding snout, narrow rostral, and uniform reddish-brown coloration, often with a dark wash. Based on the genetic and morphological differences, we redescribe T. cephalus, resurrect and elevate T. gigas to a full species and designate a neotype for this taxon, and describe three lineages as new species (T. diabolicus sp. nov., T. fortescuensis sp. nov., T. pseudopsephos sp. nov.).

Keywords: Reptilia, agamid lizard, cryptic species, Gascoyne, morphology, mtDNA, nDNA, neotype, Pilbara, taxonomy, Tympanocryptis diabolicus sp. nov., Tympanocryptis fortescuensis sp. nov., Tympanocryptis gigas, Tympanocryptis pseudopsephos sp. nov.

Doughty, Paul, Luke Kealley, Luke Shoo & Jane Melville. 2015. Revision of the Western Australian Pebble-mimic Dragon Species-Group (Tympanocryptis cephalus: Reptilia: Agamidae). Zootaxa. 4039(1): 85–117. DOI:  10.11646/zootaxa.4039.1.3

http://www.mapress.com/zootaxa/2015/f/zt04039p117.pdf

FIGURE 5. Photographs in life of Tympanocryptis cephalus species-group members (left to right, top to bottom):
  a) T. cephalus (photo—G. Harold), b) T. gigas (M. Peterson), c) T. diabolicus sp. nov. (M. Peterson), d) T. fortescuensis sp. nov. (G. Harold), e) T. pseudopsephos sp. nov. —plain form (G. Harold), f) T. pseudopsephos sp. nov.—with charcoal wash on dorsum (G. Harold).

[Cetology / Behaviour • 2015] Omura’s Whales (Balaenoptera omurai) off northwest Madagascar: Ecology, Behaviour and Conservation Needs

Omura’s whale Balaenoptera omurai

from Cerchio et al. 2015,  DOI: 10.1098/rsos.150301
courtesy of New England Aquarium || news.NEAq.org

Abstract

The Omura’s whale (Balaenoptera omurai) was described as a new species in 2003 and then soon after as an ancient lineage basal to a Bryde’s/sei whale clade. Currently known only from whaling and stranding specimens primarily from the western Pacific and eastern Indian Oceans, there exist no confirmed field observations or ecological/behavioural data. Here we present, to our knowledge, the first genetically confirmed documentation of living Omura’s whales including descriptions of basic ecology and behaviour from northwestern Madagascar. Species identification was confirmed through molecular phylogenetic analyses of biopsies collected from 18 adult animals. All individuals shared a single haplotype in a 402 bp sequence of mtDNA control region, suggesting low diversity and a potentially small population. Sightings of 44 groups indicated preference for shallow-water shelf habitat with sea surface temperature between 27.4°C and 30.2°C. Frequent observations were made of lunge feeding, possibly on zooplankton. Observations of four mothers with young calves, and recordings of a song-like vocalization probably indicate reproductive behaviour. Social organization consisted of loose aggregations of predominantly unassociated single individuals spatially and temporally clustered. Photographic recapture of a female re-sighted the following year with a young calf suggests site fidelity or a resident population. Our results demonstrate that the species is a tropical whale without segregation of feeding and breeding habitat, and is probably non-migratory; our data extend the range of this poorly studied whale into the western Indian Ocean. Exclusive range restriction to tropical waters is rare among baleen whale species, except for the various forms of Bryde’s whales and Omura’s whales. Thus, the discovery of a tractable population of Omura’s whales in the tropics presents an opportunity for understanding the ecological factors driving potential convergence of life-history patterns with the distantly related Bryde’s whales.

 KEYWORDS: Omura’s whale, Balaenoptera omurai, distribution, feeding ecology, breeding ecology, acoustic behaviour

Figure 3. Images of Madagascar Balaenoptera omurai displaying details of pigmentation and external appearance.
Five different individuals are pictured: underwater video frame captures of an adult female with calf sighted on 9 November 2013 that had just completed a feeding lunge (first row), and a lone adult on 22 October 2014 (second row), and above surface photographs from three adults sighted on 13 November 2014 (third row), 9 November 2013 (fourth row) and 12 December 2012 (fifth row). Visible features: (A) asymmetrical coloration of the lower jaw, with lightly pigmented right jaw and darkly pigmented left jaw; (B) asymmetrical coloration of the gape (inferred by inner lower lip), with lightly pigmented left gape and darkly pigmented right gape; (C) leading edge of pectoral fin white from tip to shoulder; (D) the apparent absence of lateral rostral ridges, with only faint indications detectable at some angles; (E) lightly pigmented blaze originating anterior to the eye, present only on the right side, with dark eye and ear stripe, two additional dark stripes and a light inter-stripe wash; (F) lightly pigmented chevron anterior to dorsal fin, present on both sides but asymmetrical and most prominent on right where it displays a double banded pattern; (G) highly falcate dorsal fin with gradual sloping insertion into dorsum.

 DOI: 10.1098/rsos.150301

Figure 4. An Omura’s whale subsurface lunge feeding, displaying several of the features detailed in figure 3.

 DOI: 10.1098/rsos.150301

Discussion

Prior to our discovery of Omura’s whales in Madagascan waters, there was a complete absence of field data on this species. Here we have presented evidence supporting the species identification of the small rorqual that we have observed as B. omurai, along with details on the external body appearance, and initial observations on habitat use, local distribution, feeding ecology, reproductive ecology and vocal behaviour. Notably, this represents to our knowledge, the first identified population tractable for field study. In the absence of a detailed description of the external appearance of B. omurai, it has been difficult for field biologists to confidently distinguish the species from congeners, particularly Bryde’s whales. We believe that this difficulty was primarily because of the lack of observations, and now with the detailed description we have provided it should be no more difficult to distinguish Omura’s whales in the field than it would be to distinguish fin whales from sei whales, or sei whales from Bryde’s whales, provided adequate observations and documentation.

Salvatore Cerchio, Boris Andrianantenaina, Alec Lindsay, Melinda Rekdahl, Norbert Andrianarivelo and Tahina Rasoloarijao. 2015. Omura’s Whales (Balaenoptera omurai) off northwest Madagascar: Ecology, Behaviour and Conservation Needs.
Royal Society Open Science. 2: 150301.   DOI: 10.1098/rsos.150301

[Herpetology • 2014] Hemiphyllodactylus engganoensis • A New Species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Pulau Enggano, southwestern Sumatra, Indonesia

ABSTRACT

A new species of gekkonid lizard Hemiphyllodactylus engganoensis sp. nov. from Pulau Enggano, southwestern Sumatra, Indonesia is differentiated from all other congeners by having the unique combination of a maximum SVL of 37.3 mm; six chin scales; no enlarged postmentals; five circumnasal scales; three or four scales between the supranasals; 12 supralabials; 24 dorsal scales; 15 ventral scales; a lamellar hand formula of 4554 or 4454; a lamellar foot formula of 4555; four subdigital lamellae on the first finger; four or five subdigital lamellae on the first toe; a continuous, femoroprecloacal pore series of 42; five cloacal spurs in males; no enlarged subcaudal scales; no dark postorbital stripes or striping on body; small dark blotches on dorsum; a yellowish postsacral mark bearing anteriorly projecting arms; and a pigmented caecum and gonads. Hemiphyllodactylus engganoensis sp. nov. is part of the speciose H. typus group.

Key words: Gekkonidae, Hemiphyllodactylus, Hemiphyllodactylus engganoensis sp. nov., Sumatra, new species

Hemiphyllodactylus engganoensis sp. nov.

Pulau Enggano Dwarf Gecko | Cicak Kerdil Enggano 

 L Lee Grismer, Awal Riyanto, Djoko T Iskandar and Jimmy A Mcguire. 2014. A New Species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Pulau Enggano, southwestern Sumatra, Indonesia. Zootaxa. 3821(1):485-95.

ResearchGate.net/publication/263746291_A_new_species_of_Hemiphyllodactylus_Bleeker_1860_(Squamata_Gekkonidae)_from_Pulau_Enggano_southwestern_Sumatra_Indonesia

http://ib.berkeley.edu/labs/mcguire/Grismer_et_al_2014_Hemiphyll.pdf

[Herpetology • 2014] Hemiphyllodactylus kiziriani • A New Species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Laos

ABSTRACT

A new species of the genus Hemiphyllodactylus is described from Luang Prabang Province, northern Laos. Hemiphyllodactylus kiziriani sp. nov. is distinguished from the remaining congeners by morphology, coloration, and a significant genetic divergence of greater than 20% (ND2 gene). The new species from Laos is characterized by the following features: SVL of adult males 35.1–40.1 mm, of adult females 36.3–40.8 mm; dorsal scale rows 18–27; ventral scale rows 11–15; chin scales bordering mental and first infralabial distinctly enlarged; digital lamellae formulae 3-4-4-4 (forefoot) and 4-4/5-4/5-4 (hindfoot); femoral pores 0–4, total precloacal pores 10–13 in males, 8–10 pitted precloacal scales in females; cloacal spurs present in both sexes; dorsal trunk pattern dark brown with two rows of irregular transverse bands; dark lateral head stripe distinct; upper zone of flank with a dark brown stripe; caecum and gonadal ducts unpigmented.

Keywords: Slender Gecko, karst forest, phylogeny, taxonomy, Luang Prabang Province

Truong Q. Nguyen, Andreas Botov, Minh D. Le, Liphone Nophaseud, George Zug, Michael Bonkowski and Thomas Ziegler. 2014. A New Species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Laos. Zootaxa. 3827(1):45-56. DOI: dx.doi.org/10.11646/zootaxa.3827.1.4

ResearchGate.net/publication/263544120_A_new_species_of_Hemiphyllodactylus_(Reptilia_Gekkonidae)_from_northern_Laos

[Herpetology • 2013] Integrative Taxonomy uncovers High Levels of Cryptic Species Diversity in Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) and the Description of A New Species, Hemiphyllodactylus tehtarik, from Peninsular Malaysia

Hemiphyllodactylus tehtarik 
Grismer, Wood, Anuar, Muin, Quah, McGuire, Brown, Tri & Thai, 2013

 DOI: 10.1111/zoj.12064

Conserving the planet's biodiversity is greatly handicapped, in that only a small fraction of it (∼14–75%) has been described so far. Integrative taxonomy is making significant inroads in light of this challenge by incorporating multiple data sets across a wide range of disciplines that simultaneously elucidate phylogenetic structure and delimit species-level lineages within a unified species concept. An integrative taxonomic approach to the classification of the gekkonid genus Hemiphyllodactylus Bleeker, 1860 reveals that it is far more diverse than posited by a recent taxonomic revision based solely on morphology, and that it is composed of at least 19 species, most of which are montane upland or insular endemics. Three new species (Hemiphyllodactylus dushanensis sp. nov., Hemiphyllodactylus jinpingensis sp. nov., and Hemiphyllodactylus longlingensis sp. nov.) from southern China previously considered to be subspecies of Hemiphyllodactylus yunnanensis (Boulenger, 1903) are elevated to full species status, and 10 new species-level lineages ranging from Vietnam, Laos, Thailand, Myanmar, Peninsular Malaysia, the Philippines, and Indonesia are identified. One new species, Hemiphyllodactylus tehtarik sp. nov. from Gunung Tebu, Terengganu, Peninsular Malaysia, is described herein, and is differentiated from all other species of Hemiphyllodactylus on the basis of morphology, colour pattern, and an 18.1–31.5% sequence divergence from all other congeners. Hemiphyllodactylus larutensis (Boulenger, 1900) is removed from the synonymy of H. harterti (Werner, 1900). Using an integrative taxonomic approach imbues the revised classification of Hemiphyllodactylus with more objectivity, stability, and phylogenetic history, while identifying undescribed species-level lineages in potential need of conservation.

Keywords: conservation; cryptic species; Gekkonidae; Hemiphyllodactylus tehtarik sp. nov.; Hemiphyllodactylus; integrative taxonomy; Malaysia; new species

Systematics and taxonomy

The Harterti group

Currently, there are three species of Hemiphyllodactylus recognized from Peninsular Malaysia (Grismer, 2011a): the widely distributed unisexual H. typus and the bisexual, upland species Hemiphyllodactylus harterti (Werner, 1900) from the Banjaran Bintang, and Hemiphyllodactylus titiwangsaensis Zug, 2010, from the Banjaran Titiwangsa. The latter two species are endemic to their respective mountain ranges, and make up the harterti group (Fig. 4). We report on a gravid gekkonid collected during an expedition to Gunung Tebu in the Banjaran Timur found at 600 m a.s.l. We assign this specimen to the genus Hemiphyllodactylus for having undivided, U-shaped, terminal lamellar pads on digits 2–5 of the hands and feet, free terminal phalanges that are not united with the lamellar pad, a clawless, vestigial first digit on the hands and feet, lidless eyes with vertical pupils, and for lacking a ventrolateral fringe or fold of skin extending between the axilla and the groin. However, this specimen is not referable to any known species of Hemiphyllodactylus in that it has a rust–orange, unicoloured dorsal body pattern, no white postorbital spots, and a lamellar foot formula of 3-4-5-4: characters not known from any other Hemiphyllodactylus. Additionally, it bears a unique combination of other morphological and colour pattern characters that separate it from all other known congeners. We consider this specimen to be a member of the harterti group given that it is the sister lineage of H. larutensis (see below), and describe it herein as a new species.

Tebu Mountain Slender-toed Gecko

Distribution: Hemiphyllodactylus tehtarik sp. nov. is known only from the type locality on Gunung Tebu, Terengganu, Peninsular Malaysia (Fig. 4), but is expected to range more widely throughout this portion of the Banjaran Timur.

Natural history: The holotype was seen at night on the leaf of a palm tree (Licuala sp.), approximately 1 m above the ground, along the edge of a small stream coursing through large granite boulders. During a first attempt to capture it, the lizard dropped to the ground and escaped, but was captured later that night on a different leaf of the same tree. The holotype is an adult female carrying two eggs, indicating that the reproductive season of this species extends at least through August. From only a single female, it cannot be determined if H. tehtarik sp. nov. is unisexual or bisexual.

Etymology: This species is named after a traditional Malaysian tea, Teh Tarik, which bears the rich orangish coloration of the holotype.

Discussion

It is a well-established fact that the world's biodiversity is woefully underestimated (Mora et al., 2011), and that most species with broad distributions across environmentally heterogeneous regions are rarely a single species, but rather complexes of generally cryptic species. This is especially true in tropical regions such as Southeast Asia, the dynamic and complex environmental history of which (Hall, 1998, 2001, 2002, 2012; Woodruff, 2003, 2010; Bird, Taylor & Hunt, 2005; Outlaw & Volker, 2008; Reddy, 2008; Cannon, Morley & Bush, 2009; Cannon, 2012; Gower et al., 2012; Morley, 2012; Richardson, Costion & Muellner, 2012) has contributed greatly to the phylogenetic and phylogeographic structuring of a number of wide-ranging clades thought previously to be single species (e.g. Matsui et al., 2005; Stuart, 2006; Inger, Stuart & Iskandar, 2009; Wood et al., 2009, 2010; Grismer et al., 2010b, 2012a; Malhotra et al., 2011; Brown et al., 2012; Johnson et al., 2012; Nishikawa et al., 2012; Loredo et al., 2013). The infusion of a molecular phylogenetic data set into the morphology-based classification of the wide-ranging genus Hemiphyllodactylus not only contributes to this growing body of knowledge, but highlights the inherent tendency of morphological data sets to underestimate biodiversity in taxonomic revisions of widespread groups. Zug's (2010) characterization of Hemiphyllodactylus as a ‘low diversity taxon’ is emblematic of this problem, in that this group's diversity was underestimated conservatively by as much as 113–162%. The molecular phylogeny of Hemiphyllodactylus (Fig. 1) facilitated an independent test of the morphology-based taxonomy (Zug, 2010), identified a minimum of ten (perhaps as many as 13) additional lineages for consideration of species-level designation, and resulted in a less subjective and more stable classification upon which others can build. Some (i.e. Isaac, Mallet & Mace, 2004) posit that studies such as these: (1) result in taxonomic inflation because they tend to use lineage-based species concepts as opposed to a more ‘stable’ biological species concept; (2) impede macroevolutionary studies because they generate asymmetric taxonomies across groups, making them incomparable; and (3) generate unnecessary challenges for conservation efforts. We argue that rather than inflating taxonomies, an integrative approach based on a unified lineage concept (de Queiroz, 1998) actually reveals the true structure of biodiversity, and will ultimately enhance conservation efforts by bringing to the forefront the entities (i.e. species, management units, etc.) in actual need of conserving. The resulting taxonomies are not only more species rich, but are more stable because they are inextricably bound to methodologically and conceptually less biased molecular data sets, and are delivered from the pluralistic outdated biological species concept that is difficult or impossible to apply to allopatric lineages, and often misrepresents the history (e.g. see Frost & Kluge, 1994).

Recent taxonomic work across the broad spectrum of life is revealing that nature is considerably more diverse than a polytypic species concept (Mayr, 1963) would lead us to believe. And if we are to forward a sincere effort in an attempt to stem the biodiversity crisis, we need to know upon what to focus our efforts. The very broad distribution (from the Mascarene Islands to Hawaii) and generally conserved morphology of Hemiphyllodactylus illustrates the limits of a morphological taxonomy in the delimitation of species boundaries for such groups. Considerable work on this group is still necessary before we gain a clear understanding of its diversity. The genetics of many of the known populations from Vietnam, Laos, Cambodia, Thailand, and throughout the islands of Indonesia, for example, remain unsampled. As additional material becomes available, we will use the phylogentic taxonomy hypothesized here as a foundation upon which to build future taxonomies. Infusing this taxonomy with molecular estimates of phylogeny allows for a more fine-grained analysis and greater potential for discovering species-level lineages and distinct management units that may be of special concern and in need of conservation (Hodkinson et al., 2012).

L. Lee Grismer, Perry L. Wood Jr, Shahrul Anuar, Mohd Abdul Muin, Evan S. H. Quah, Jimmy A. McGuire, Rafe M. Brown, Ngo Van Tri and Pham Hong Thai. 2013. Integrative Taxonomy uncovers High Levels of Cryptic Species Diversity in Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) and the Description of A New Species from Peninsular Malaysia. Zoological Journal of the Linnean Society. 169(4); 849–880. DOI: 10.1111/zoj.12064

http://ib.berkeley.edu/labs/mcguire/Grismer_et_al_Hemiphyllo_2013.pdf

[Herpetology • 2013] Hemiphyllodactylus zugi • A New Species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Vietnam

Hemiphyllodactylus zugi

Thạch sùng dẹp zug | Zug’s  Slender Gecko 

Nguyen, Lehmann, Le, Duong, Bonkowski & Ziegler, 2013

Abstract

We describe a new species of the genus Hemiphyllodactylus on the basis of four specimens from Cao Bang Province, northern Vietnam. Hemiphyllodactylus zugi sp. nov. is distinguished from the remaining congeners by a combination of the following characters: a bisexual taxon; average SVL of adult males 41 mm, of adult female 46.6 mm; chin scales bordering mental and first infralabial distinctly enlarged; digital lamellae formulae 3-4-4-4 (forefoot) and 4-5-5-5 (hindfoot); femoral and precloacal pore series continuous, 18–21 in total in males, absent in female; cloacal spur single in males; dorsal trunk pattern of dark brown irregular transverse bands; dark lateral head stripe indistinct; upper zone of flank with a series of large light spots, edged above and below in dark grey; caecum and gonadal ducts unpigmented.

Keywords: Slender Gecko, karst forest, phylogeny, taxonomy, Cao Bang Province, Ha Lang District

Hemiphyllodactylus zugi 

Etymology. We name the new species in honour of Professor Dr. George Zug, National Museum of Natural History,  Smithsonian  Institution  (Washington  DC,  USA),  in  recognition  of  his  contribution  to  herpetological  research  in  Southeast  Asia,  particularly  the  genus  Hemiphyllodactylus.  As  common  names  we  suggest  Zug’s  Slender Gecko (English), Zugs Halbblattfingergecko (German), and Thạch sùng dẹp zug (Vietnamese).

Natural history. Hemiphyllodactylus zugi inhabits disturbed secondary limestone forests near a residential area at elevations between 435–601 m. Specimens were found at night on a wooden fence, tree bark and leaves near cave entrances or on a limestone boulder near a forest path, ca. 0.5–1.5 m above the ground. 

Distribution. The species is currently known only from Cao Bang Province in northern Vietnam (Fig. 4). This species is expected to be found in the contiguous limestone forests in Guangxi Province, China as in the case of Gekko adleri Nguyen, Wang, Yang, Lehmann, Le, Ziegler & Bonkowski, a recently described species from the border region between China and Vietnam (Nguyen et al. 2013).  

Truong Quang Nguyen, Tanja Lehmann, Minh Duc Le, Ha Thuy Duong, Michael Bonkowski & Thomas Ziegler. 2013. A New Species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Vietnam. Zootaxa. 3736 (1): 089–098

http://biotaxa.org/Zootaxa/article/view/zootaxa.3736.1.5

ResearchGate.net/publication/258374068_A_new_species_of_Hemiphyllodactylus_(Reptilia_Gekkonidae)_from_northern_Vietnam

[Botany • 2015] Besleria macropoda (Gesneriaceae): Lectotypification, Distribution, Functional Epiphylly and Discordant Fruit Morphology of A Rare Costa Rican Endemic

FIGURE 2. Besleria macropoda, epiphyllous position of flowers and fruits.
A. Shoot. B. Inflorescence with open flowers. C. Infructescence showing fruits with split and reflexed carpel walls exposing red placentae and seeds. D. Sulcate petiole and leaf base clasping the peduncle above the sunken midrib.

(Photos by A. Berger; voucher: Berger & Etl 1443.).DOI: 10.11646/phytotaxa.233.2.2

Abstract

Besleria macropoda, a rare and poorly known gesneriad endemic to Costa Rica, was recently collected for the first time on the southern slopes of the Fila Costeña (Puntarenas Province, SE Costa Rica). The collection considerably widens the geographic distribution to the southeastern part of Costa Rica. Moreover, the following unique characters not previously addressed in the literature were observed and are documented here: (1) The elongate peduncles of the inflorescences are clamped in a channel formed by the sunken midrib of the leaf, rendering the flowers and fruits positioned in the center of the leaf blade. The epiphyllous appearance of the inflorescence on the leaf surface enhances contrasting colors that may aid the pollination and/or fruit dispersal. (2) The fruits split open irregularly, with the fleshy carpel lobes becoming reflexed. This fruit dehiscence deviates from the indehiscent berries that typically characterize Besleria. This results in displaying a globose head of red placental tissue covered by tiny, red seeds. A preliminary survey of Besleria fruits suggests that this peculiar fruit type is present in at least 15 species representing almost 8% of the genus. Fruit morphology of Besleria is therefore less uniform than previously recognized and the “indehiscent berry” can no longer serve as a distinctive generic character of Besleria, which necessitates consideration in floras and identification keys. In addition, a list of herbarium specimens, lectotypification, a distribution map, IUCN red list assessment and an amended key to diagnose Besleria relative to Gasteranthus are provided.

Keywords: Besleria, Gesneriaceae, fruit morphology, epiphylly, Eudicots, Costa Rica

Conclusions

The recent collection of Besleria macropoda widens the known distribution considerably and highlights two unique characters not previously addressed. The inflorescences/infructescences are functionally epiphyllous possibly aiding in pollination and/or fruit dispersal. Fruit dehiscence deviates from indehiscent berries that typically characterize Besleria: the fruits appear ruptured and split open irregularly, with the fleshy carpel lobes becoming reflexed. This results in displaying a globose head of red placental tissue embedded with tiny, red seeds. In addition to B. macropoda, this specific fruit type was found in 14 other species of the genus. Fruit morphology of Besleria is thus less uniform than previously understood, and the “indehiscent berry” can no longer serve as a distinctive generic character of Besleria, which necessitates consideration in generic keys such as in regional floras. However, further fieldwork is necessary to fully understand the distribution, phylogenetic context, and functional implications of this particular fruit type and fruit dehiscence.

Andreas Berger, John J. Clark and Anton Weber. 2015. Besleria macropoda (Gesneriaceae): Lectotypification, Distribution, Functional Epiphylly and Discordant Fruit Morphology of  A Rare Costa Rican Endemic. Phytotaxa. 233(2): 139–152. DOI: 10.11646/phytotaxa.233.2.2

[Crustacea • 2015] Aletheiana tenella • A New Genus and New Species of Freshwater hymenosomatid Crab (Decapoda: Brachyura) from Central Sulawesi, Indonesia

Aletheiana tenella

Ng, & Lukhaup, 2015

FIGURE 1. Aletheiana tenella gen. et sp. nov.. A, cascades at Puawu River, central Sulawesi, B, submerged roots at the riverbank of Puawu River where the specimens were collected; C–F, colour in life.

DOI: 10.11646/zootaxa.4039.1.4

Abstract 

A new genus and new species of free-living hymenosomatid crab, Aletheiana tenella, is described from Central Sulawesi, Indonesia. The two known Sulawesi hymenosomatid species, Cancrocaeca xenomorpha Ng, 1991, and Sulaplax ensifer Naruse, Ng & Guinot, 2008, are both from cave habitats. Aletheiana gen. nov. is most similar to Neorhynchoplax Sakai, 1938 (from freshwater and intertidal habitats in the Indo-West Pacific), and Sulaplax, but can be distinguished by its front possessing only one subventral rostral lobe, the base of the antenna is positioned between the base of the ocular peduncle and antennular fossa, the posterior margin of the epistome has two low, rounded median lobes, the merus of the third maxilliped is elongated, the ambulatory dactylus has a prominent subdistal spine, the cutting edges of the chela are armed with distinct teeth proximally, and the male abdomen is slender and elongate with the telson linguiform.

Key words: Hymenosomatidae, Aletheiana tenella, new genus, new species, taxonomy, Sulawesi

Etymology. “Aletheia” is an ancient-Greek word for disclosure and a state of not being hidden. The latinised name is used here as a genus, alluding to the present discovery of the first free-living hymenosomatid crab from Sulawesi. Gender of genus feminine. The name is derived from the Latin “tenellus” for delicate; alluding to the small size and slender appendages of the species.

Biology. Aletheiana tenella gen. et sp. nov. was found among the submerged fibrous roots of trees at the slow flowing parts of the riverbank near cascades along the Puawu River, in Central Sulawesi (Fig. 1A, B). The crabs were observed at the slow flowing parts of the riverbank among roots near cascades of the Puawu River. The water at the time of collection was muddy and had a temperature of 22°C. In the faster flowing parts of the river, no crabs were found. This is a typical habitat for many free-living hymenosomatids (e.g., see Naruse & Ng 2007). A new species of Caridina, C. boehmei (Atyidae) was also recently described from the type locality of Aletheiana tenella gen. et sp. nov. by Klotz & von Rintelen (2013).

Ng, Peter K. L. and Christian Lukhaup. 2015. Aletheiana tenella, A New Genus and New Species of Freshwater hymenosomatid Crab (Crustacea: Decapoda: Brachyura) from Central Sulawesi, Indonesia. Zootaxa. 4039(1): 118–128. DOI: 10.11646/zootaxa.4039.1.4

facebook.com/photo.php?fbid=10153359002238198
http://freepdfhosting.com/1aed2ea4e4.pdf

[Herpetology • 2015] Malayodracon gen. nov. • The Systematic Status of Gonocephalus robinsonii Boulenger, 1908 (Agamidae: Draconinae) | กิ้งก่าดงเขาสูง

Malayodracon gen. nov. 

Denzer, Manthey, Mahlow & Böhme, 2015

กิ้งก่าดงเขาสูง | Malayodracon robinsonii (Boulenger, 1908)

FIGURE 5. Malayodracon robinsonii from Tanah Rata, Cameron Highlands, West Malaysia.

Photo: U. Manthey. 
DOI: 10.11646/zootaxa.4039.1.5 || ResearchGate.net

Abstract

The generic assignment of the draconine lizard Gonocephalus robinsonii from the highlands of West-Malaysia has been uncertain since the original description. Here we present a study based on morphology, previously published karyotype data and molecular phylogenetics using 16S rRNA sequences to evaluate the systematic status of G. robinsonii. As a result we describe Malayodracon gen. nov. to accommodate the species.

Keywords: Reptilia, Gonocephalus, phylogeny, cranial morphology, Malayodracon gen. nov., Malayodracon robinsonii comb. nov., Dendragama boulengeri

Malayodracon gen. nov. 

Type species: Gonyocephalus robinsonii Boulenger, 1908;
Type locality: Gunong (=Gunung / Mt.) Tahan (5,200 ft / 1,585 m), Pahang, Malaysia; the holotype (by monotypy) is deposited in the British Museum Natural History BMNH 1946.8.14.81 (previously BMNH 1906.2.28.8); collected by H.C. Robinson. 

Taxon — Malayodracon robinsonii (Boulenger, 1908)

Chresonymy

Gonyocephalus robinsonii — Boulenger 1908: 65; Boulenger 1912: 67

Gonyocephalus robinsoni — Smith 1922: 269

Gonocephalus robinsoni — Smith 1930: 24; Smirnova 2003: 128

Goniocephalus robinsoni — Smith 1935: 133; Sly 1976: 156; Bourret 1947 “2009”: 211

Gonocephalus robinsonii — Smedley 1931: 110; Wermuth 1967: 61; Moody 1980: 299; Denzer & Manthey 1991: 312; Manthey & Denzer 1992a: 16; Manthey & Schuster 1992: 65; Manthey & Grossmann 1997: 189; Chan-ard et al. 1999: 102; Diong et al. 2000: 73; Honda et al. 2002; Denzer & Manthey 2009: 256; Grismer 2011: 258; Pyron et al. 2013

(Gonocephalus inc. sed.) robinsonii — Manthey 2010: 46

Distribution. Malayodracon robinsonii is restricted in its distribution from mid to high altitude areas of the peninsular Malaysian highlands (approx. 600–1500 m asl) inhabiting submontane and montane forests. M. robinsonii shares this isolated distribution with several other reptile species that are restricted to the Cameron Highlands or the central mountain ranges of Malaysia (Fraser and Larut Hills; Genting Highlands) such as Trimeresurus nebularis Vogel, David & Pauwels, 2004, Hebius sanguineus (Smedley, 1931), Macrocalamus tweediei Lim, 1963 and Collorhabdium williamsoni Smedley, 1931. Endemism on genus and species level is comparatively high in this geographical area.
Malayodracon robinsonii has been reported from Genting Highlands (Ulu Kali), Cameron Highlands (Tanah Rata, Gunung Brinchang, Gunung Beremban, Gunung Jesar) and further north from its type locality Gunung Tahan.

Etymology. The name Malayodracon was chosen to express that the type and currently only known species of the genus is restricted to Malaysia (latinized malaya, male form malayo owing to the gender of the ending -dracon) and constitutes a genus belonging to the subfamily Draconinae (gr. drakon / δράκων; a serpent in Greek mythology; latinized dracon = engl. dragon). 

Denzer, Wolfgang, Ulrich Manthey, Kristin Mahlow & Wolfgang Böhme. 2015. The Systematic Status of Gonocephalus robinsonii Boulenger, 1908 (Squamata: Agamidae: Draconinae). Zootaxa. 4039(1): 129–144. DOI: 10.11646/zootaxa.4039.1.5

ResearchGate.net/publication/283452998_The_systematic_status_of_Gonocephalus_robinsonii_Boulenger_1908_(Squamata_Agamidae_Draconinae)

[Herpetology • 2015] Boophis boppa • A New Species of Bright-eyed Treefrog (Mantellidae) from Madagascar, with comments on Call Evolution and Patterns of Syntopy in the Boophis ankaratra complex

Boophis boppa Hutter & Lambert, 2015

DOI:  10.11646/zootaxa.4034.3.6

ABSTRACT

We describe a new species of Boophis treefrog from Ranomafana National Park in the southern central east of Madagascar. This region has remarkably high anuran diversity, and along with neighbouring sites, hosts more than 35 Boophis species. Boophis boppa sp. nov. is part of the B. ankaratra sub-clade (herein named the B. ankaratra complex), previously identified within the monophyletic B. albipunctatus species group. It occurs sympatrically with two other species of the complex (B. ankaratra and B. schuboeae). Morphological differentiation of species within the B. ankaratra clade remains elusive, but species are well characterized by distinct advertisement calls, with B. boppa having the longest note duration and inter-note intervals when compared to closely related species. Furthermore, it has moderate differentiation in mitochondrial DNA, with pairwise distances of 1.9–3.7% to all other species in sequences of the mitochondrial 16S rRNA marker. Additional evidence is given by the lack of haplotype sharing with related species for the nuclear exon DNAH-3. All examples of syntopic occurrence in this complex involve species with strongly different advertisement calls, while allopatric species have more similar calls. Such a pattern might result from adaptive call co-evolution but could also be the result of non-adaptive processes. Thorough clarification of the systematics of the B. ankaratra sub-clade is required, and we outline future directions for both bioacoustic and genetic research.

Keywords: Amphibia, Anura, Ranomafana National Park, new species, reinforcement, Boophis boppa, taxonomy

Boophis boppa sp. nov.

Suggested common English name. Boppa’s Bright-eyed treefrog

Suggested common Malagasy name. Fity maso hazo Sahona ny Boppa

Etymology. This species in named in honor of Nicholas Jay Pritzker, devoted grandfather, father and husband, brilliant businessman, philanthropist, amateur scientist, committed conservationist; and board member, supporter and long-time friend of Conservation International. The name ‘Boppa’ is an affectionate nickname used by his children and grandchildren. This dedication is courtesy of his youngest son Isaac, who has generously supported amphibian research in Madagascar, including this study and future taxonomic research as well. The species name is used as an invariable noun in apposition to the genus name. 

Natural history: Boophis boppa was locally abundant but thus far only found in undisturbed, primary forests along fast moving streams and was occasionally found along slow flowing tributaries adjacent to fast streams. Males of the species typically were calling at night from the surfaces of vegetation less than three meters in height (Fig. 12A–B). Females of B. boppa were rarely encountered and were only observed while in amplexus (Fig. 12C). We also confirm that Boophis boppa is syntopic with Boophis ankaratra at Maharira and that the two species can be found calling less than a meter apart. This is also consistent with Andreone et al. (2007), which found B. boppa (i.e. Boophis aff. ankaratra) and B. ankaratra to occur sympatrically. Other syntopic species of Boophis at Maharira include: B. madagascariensis, B. majori, B. aff. marojezensis, B. aff. picturatus, B. popi, and B. reticulatus. 

 Carl R. Hutter and Shea M Lambert. 2015. A New Species of Bright-eyed Treefrog (Mantellidae) from Madagascar, with comments on Call Evolution and Patterns of Syntopy in the Boophis ankaratra complex. ZOOTAXA. 4034(3):531-555.  DOI:  10.11646/zootaxa.4034.3.6

ResearchGate.net/publication/283318430_A_new_species_of_bright-eyed_treefrog_(Mantellidae)_from_Madagascar_with_comments_on_call_evolution_and_patterns_of_syntopy_in_the_Boophis_ankaratra_complex

[Crustacea • 2015] Actaea grimaldii • A New Species of Reef Crab (Brachyura, Xanthidae) from Papua New Guinea

Actaea grimaldii Ng & Bouchet, 2015

DOI: 10.5852/ejt.2015.140

Abstract

A new species of xanthid crab, Actaea grimaldii, is described from the coral reefs of Papua New Guinea. This species has a distinctive red and white coloration and is closest to Actaea spinosissima Borradaile, 1902, from the Indian Ocean. However, the new species can be distinguished by the arrangement of spines on the carapace, chelipeds and ambulatory legs, and the structure of the male gonopods. Actaea grimaldii sp. nov. has also been confused with A. polyacantha (Heller, 1861), but differs markedly in the carapace armature.

Keywords: Crustacea, Xanthidae, Papua New Guinea, Actaea, new species.

Infraorder Brachyura Latreille, 1802

Superfamily Xanthoidea MacLeay, 1838

Family Xanthidae MacLeay, 1838

Subfamily Actaeinae Alcock, 1898

Genus Actaea De Haan, 1833

Actaea grimaldii sp. nov.

urn:lsid:zoobank.org:act:A736C0FE-D2B4-4256-AB85-4370386B0231

Figs 1–5

Actaea peronii var. squamosa – Calman 1900: 10 (not Actaea squamosa Henderson, 1893).

Actaea spinosissima – Odhner 1925: 59 (part). — Serène 1984: 115, pl. 14F. — Davie 2002: 511 (not

Actaea spinosissima Borradaile, 1902).

Etymology: The species name is in honour of His Serene Highness Albert II, Prince of Monaco, patron of the PAPUA NIUGINI Expedition and several other biodiversity expeditions of the “Our Planet Reviewed” programme conducted by MNHN and Pro-Natura International. The red and white colour pattern of the new species also alludes to the colours associated with the House of Grimaldi.

Distribution: This species is known for certain from the Moluccas and Papua New Guinea, and is probably also present in Australia.

Peter K.L. Ng and Philippe Bouchet. 2015. Actaea grimaldii, A New Species of Reef Crab from Papua New Guinea (Crustacea, Brachyura, Xanthidae). European Journal of Taxonomy. 140: 1–18. DOI: 10.5852/ejt.2015.140

http://www.europeanjournaloftaxonomy.eu/index.php/ejt/article/view/262/500

http://lkcnhm.net/2015/10/16/actaea-grimaldii-a-crab-of-royal-status/