[Ichthyology • 2021] A Taxonomic Review of the Family Trachipteridae (Acanthomorpha: Lampridiformes), with An Emphasis on Taxa Distributed in the western Pacific Ocean

Zu sp. 

in Martin & Hilton, 2021. 

DOI: 10.11646/ZOOTAXA.5039.3.1

photographed in Nusa Penida, Indonesia by Helen Mitchell

Abstract

The family Trachipteridae—the Ribbonfishes, Dealfishes, and their relatives—has a circumglobal distribution, with at least 10 species in three genera (Zu Walters & Fitch 1960, Desmodema Walters & Fitch 1960, and Trachipterus Goüan 1770) that are characterized by elongate, extremely laterally compressed bodies, large eyes, absence of ribs, spines on lateral-line scales, greatly protrusible mouths, and a lack of pelvic fins in adults. They are also known for the drastic morphological changes that occur during ontogeny. Trachipterids are poorly represented in collections due to the fragile nature of their bodies. Most studies of the Trachipteridae have been limited by the numbers, developmental stages, and the completeness of the specimens that were examined. Along with the lack of available material, incomplete and conflicting character information compounds the taxonomic confusion of Trachipteridae. Despite the body of regional revisions that have examined trachipterid taxonomy, none have synthesized a suite of morphological characters across ontogeny. The goals of this paper are to (1) revise the family Trachipteridae, (2) revise the genera Trachipterus, Zu, and Desmodema, including information regarding ontogeny and biogeography, and 3) address the alpha taxonomy of Zu, Desmodema, and Trachipterus from the western Pacific Ocean. We recognize possibly five species of Trachipterus as being present in the western Pacific, as well as two species of both Zu and Desmodema. Despite additions to the specimen base that allows refinement of taxonomy and diagnoses, there are still large knowledge gaps associated with the taxonomic review of Trachipteridae. These reflect incomplete understanding of geographic distribution of taxa which may mask unrecognized taxonomic variability. The genus Trachipterus specifically remains problematic and will require greater detailed global study. Early life history stages remain unknown for several taxa which hinders full interpretation of ontogenetic transitions. Protracted transitions, some of which are clarified here, further confuse stage-based diagnoses and must be considered in future analyses of this family.

 

Keywords: Pisces, Ribbonfishes, Trachipterus, Desmodema, Zu, ontogenetic characters 

Order Lampridiformes Goodrich 1909

Family Trachipteridae Swainson 1839

Ribbonfishes, Dealfishes

Trachipterus Goüan 1770

Putatively valid taxa in the western Pacific Ocean 

1. Trachipterus trachypterus (Gmelin 1789). Holotype: No types known. Mediterranean. 

2. Trachipterus altivelis Kner 1859. Syntype: NMW 22046. Valparaiso, Chile. Unavailable for examination in current study.

3. Trachipterus arawatae Clarke 1881. Holotype NMNZ P.1008, 51 cm SL. Hominy Cove, Jackson’s Bay New Zealand. Examined.

4. Trachipterus jacksonensis (Ramsay 1881). Syntype: AMS A.9114, 1408 mm, caudal missing (736 mm SV). Manly Beach, Port Jackson, New South Wales, Australia. Examined.

5.  Trachipterus ishikawae Jordan & Snyder 1901. Holotype: NSMT 589, 1250 mm SL. Off the mouth of Tokyo Bay, between Misaki and Boshu. Examined.

Zu sp. Live specimen (not collected) photographed in Mallorca, Spain.

photo: Michael Makowiecki.

DOI: 10.11646/ZOOTAXA.5039.3.1

Zu sp. Live specimen (not collected) photographed in Nusa Penida, Indonesia.

photo: Helen Mitchell.

DOI: 10.11646/ZOOTAXA.5039.3.1

Zu Walters & Fitch 1960

Zu cristatus (Bonelli 1820)

Zu elongatus Heemstra & Kannemeyer 1984

Desmodema Walters & Fitch 1960

Desmodema polystictum (Ogilby 1898)

Desmodema lorum Rosenblatt & Butler 1977

Conclusions: 

Despite advances in the understanding of Trachipteridae resulting from tremendous additions to the specimen base, which have allowed for the refinement of the taxonomy and diagnoses within the family, there are still large knowl-edge gaps associated with the taxonomy of the family. These reflect incomplete understanding of geographic dis-tribution of taxa that may mask unrecognized taxonomic variability. The genus Trachipterus specifically remains problematic and will require greater detailed global study. Early life history stages remain unknown for several taxa, which hinders full interpretation of ontogenetic transitions. Protracted transitions, some of which are clarified here, further confuse stage-based diagnoses and must be considered in future analyses of this family.

Jennifer M. Martin and Eric J. Hilton. 2021. A Taxonomic Review of the Family Trachipteridae (Acanthomorpha: Lampridiformes), with An Emphasis on Taxa Distributed in the western Pacific Ocean. Zootaxa. 5039(3); 301-351. DOI: 10.11646/ZOOTAXA.5039.3.1

[PaleoOrnithology • 2021] Archaehierax sylvestris • An Exceptional Partial Skeleton of A New Basal Raptor (Accipitriformes: Accipitridae) from the late Oligocene Namba Formation, South Australia

Archaehierax sylvestris

Mather, Lee, Camens & Worthy, 2021

DOI: 10.1080/08912963.2021.1966777

 

ABSTRACT

The Australian pre-Pleistocene fossil record of Accipitridae (eagles, hawks, old-world vultures) comprises one latest Oligocene or early Miocene and one middle Miocene species, each represented by partial bones. Globally, most fossil accipitrids are based on single bones. The recent discovery of an older and considerably more complete accipitrid from late Oligocene sediments in Australia is therefore significant. It is derived from the Pinpa Local Fauna from the Namba Formation at Lake Pinpa, South Australia (~26–24 Ma). The fossil, described as Archaehierax sylvestris gen. et sp. nov., represents a raptor that was larger than the black-breasted buzzard Hamirostra melanosternon but smaller and more gracile than the wedge-tailed eagle Aquila audax. Comprehensive morphological and molecular phylogenetic analyses resolved Archaehierax as a basal accipitrid, not closely related to any living subfamily and perhaps the sister taxon to all other accipitrids exclusive of elanines. Relatively short wings similar to species of Spizaetus and Spilornis suggest it was adapted for flight within enclosed forests. Additional accipitrid fossils from the Namba Formation, a distal femur and a distal humerus, are incomparable with the holotype of A. sylvestris; they may represent distinct species or smaller individuals of the new taxon.  

KEYWORDS: Lake Pinpa, Accipitriformes, Cenozoic fossil birds, accipitrid evolution, Australia

Skeleton of an osprey by Eyton (1867) with shaded sections indicating which bones are preserved in the fossil of Archaehierax sylvestris.

 A comparison of the prepared fossil tarsometatarsus (foot bone) and a hypothesised silhouette of Archaehierax sylvestris (left) compared to the wedge-tailed eagle Aquila audax (right).

The scale bar is 10 mm long.

Systematic palaeontology

Class Aves Linnaeus, 1758

Order Accipitriformes Vieillot 1816 Family Accipitridae Vigors, 1824

Subfamily: Archaehieraxinae subfam. nov.

Type genus: Archaehierax gen. nov.

Remarks: 

The fossil is identified as an accipitrid due to the following combination of characters: Skull – Rostrum deep and narrow, with hooked tip and a large, broad nasal aperture; Tibiotarsus – Pons supratendineus ossified, aligned steeply transversely, medially placed, with unbranched canalis tendinosus, and distal condyles much wider than craniocaudally deep; Tarsometatarsus – Robust, with monosulcate hypotarsus, the lateral and medial hypotarsal crests widely separated and trochleae metatarsorum splayed both medially and laterally, and dorsally arched in distal view; Foot – Four digits with raptorial unguals, those of digits 1 and 2 relatively large; Digit IV – phalanges 2 and 3 are very short compared to phalanx 4.

The fossil can be excluded from Falconiformes (Falconidae) and the other families of Accipitriformes (Cathartidae, Sagittariidae, Pandionidae) by the morphology of the tarsometatarsal hypotarsus cristae and sulcus. The cristae are fused or partially fused together to enclose the sulcus in Cathartidae, Sagittariidae, and Pandionidae, while in Falconidae the medial crista is connected to the shaft by a ridge that extends two-thirds of its length, features that are absent in the fossil.

Diagnosis: 

Accipitrids in which the following autapomorphic features are found: the pila medialis of the sternum dorsally separates two deep pneumatic fossae, the humerus has the caput humeri only slightly elevated proximally past the tuberculum ventralis, the tip of the processus procoracoideus of the coracoid sharply curves inwards ventrally towards the medial face of the bone, the tibiotarsus has the lateral/distal retinaculum scar in a deep fossa, the tarsometatarsus is relatively elongate with narrow trochleae metatarsorum that are separated by wide incisurae, and the incisura for the m. flexor hallucis brevis tendon is large, distinct, and extends distal to the fossa metatarsi I. In addition to this, the following features occur: the rostral tip of the rostrum is hooked below the tomial margin at a relatively shallow 30–40° angle, the quadrate has a deep, distinct foramen pneumaticum caudomediale, and the sternum has the apex carinae displaced caudally from the base of the spina externa.,

Genus Archaehierax Mather, Lee, Camens and Worthy gen. nov.

 

Etymology: Archaehierax is derived from the Greek words ‘archaios’, meaning ancient, and ‘hierax’, meaning hawk. Gender masculine.

Archaehierax sylvestris Mather, Lee, Camens and Worthy gen. et sp. nov.

Etymology: The species name ‘sylvestris’ is derived from the Greek world ‘sylvas’, meaning forest, and the Latin suffix ‘-estris’, meaning ‘belonging to’.

Ellen K. Mather, Michael S. Y. Lee, Aaron B. Camens and Trevor H. Worthy. 2021. An Exceptional Partial Skeleton of A New Basal Raptor (Aves: Accipitridae) from the late Oligocene Namba Formation, South Australia. Historical Biology: An International Journal of Paleobiology. DOI: 10.1080/08912963.2021.1966777

25-million-year-old ancient eagle ruled the roost in Australia

63 bone fossil finding in desert is one of the “best preserved” eagles ever and a very rare discovery

 eurekalert.org/news-releases/929172

[Entomology • 2021] Integrative Taxonomy reveals Remarkable Diversity in Australian Protomiltogramma (Diptera: Sarcophagidae)

Protomiltogramma sp. 

Johnston, Wallman, Szpila & Pape, 2021

DOI: 10.11646/zootaxa.5043.1.1 

 twitter.com/nature_nik 

 

Abstract

Protomiltogramma Townsend is the largest and most diverse genus of miltogrammine flesh flies in Australia. However, no comprehensive taxonomic work had been completed on the Australian members of this genus in almost a century. This study presents the first taxonomic revision of all Australian species of Protomiltogramma (Sarcophagidae: Miltogramminae), completed using an integrative approach combining molecular and morphological data. Eight new species endemic to Australia are described: Protomiltogramma dalbiensis sp. n., P. grandis sp. n., P. incana sp. n., P. kapnos sp. n., P. nigrisensa sp. n., P. popularis sp. n., P. rubra sp. n. and P. subtilis sp. n. In addition, P. australis Malloch, 1930 is synonymised with P. cincta Townsend, 1916, syn. n. and P. mallochi Verves, 1987 is synonymised with P. laticeps Malloch, 1930, syn. n. Molecular phylogenetics is used to place the Australian Protomiltogramma among the miltogrammine genera of the world.

Keywords: Australia, DNA barcoding, flesh flies, Miltogramminae, phylogenetics, Diptera

 

 Nikolas P. Johnston, James F. Wallman, Krzysztof Szpila and Thomas Pape. 2021. Integrative Taxonomy reveals Remarkable Diversity in Australian Protomiltogramma (Diptera: Sarcophagidae). Zootaxa. 5043(1); 1-104. DOI: 10.11646/zootaxa.5043.1.1

  twitter.com/nature_nik/status/1442340025413169152

[Paleontology • 2022] Eptalofosuchus viridi • First Upper Cretaceous notosuchians (Crocodyliformes) from the Uberaba Formation (Bauru Group), southeastern Brazil: Enhancing Crocodyliform Diversity

 Eptalofosuchus viridi

Marinho, Martinelli, Basilici, Soares, Marconato, Ribeiro & Iori, 2022

 DOI: 10.1016/j.cretres.2021.105000 

Illustration by Júlia d'Oliveira

 twitter.com/tupandactylus 

Highlights: 

• Here we describe the first crocodyliforms fossils from the Uberaba Formation.

• The material comprises of isolated teeth and a fragmentary mandible of notosuchians.

• The isolated teeth represent Bauruchidae and a probable longisrotrine Peirosauridae.

• The fragmentary mandible was identified as a new taxon, Eptalofosuchus viridi gen et sp. nov..

Abstract

Notosuchians are the most diverse group and are frequently found in rocks of the Upper Cretaceous Bauru Group of southeastern Brazil. These crocodyliforms occupied several different terrestrial ecological niches, ranging from small to large bodied forms, including hypercarnivore to strictly herbivore animals. Most of this diversity comes from the Adamantina and Serra da Galga formations, where baurusuchids, peirosaurids and the so called “advanced notosuchians” are common occurrences. In this context, the Serra da Galga Formation (formerly Serra da Galga Member of the Marília Formation), that crops out at the Uberaba region, stands out as one of the most prolific Upper Cretaceous units in Brazil. The Uberaba Formation is another unit present in this area, underlying the Serra da Galga Formation, but to date, its crocodyliform diversity is virtually unknown. Here we describe the first notosuchians remains from the Uberaba Formation, and the first new taxon from this unit, Eptalofosuchus viridi gen. et sp. nov., based on a partial dentary with some teeth of an “advanced notosuchian”. The other remains include isolated teeth of a large baurusuchid and a possible longirostrine peirosaurid, suggesting that the Uberaba Formation shared a similar notosuchian fauna with that present in the Adamantina and São José do Rio Preto formations.

Keywords: Notosuchia, Baurusuchidae, Peirosauridae, Bauru Basin, Minas Gerais State

Archosauria Cope, 1869

Crocodylomorpha Walker, 1970 (sensu Clark, 1986)

Crocodyliformes Hay, 1930 (sensu Clark, 1986)

Mesoeucrocodylia Whetstone and Whybrow, 1983

Notosuchia Gasparini, 1971 (sensu Pol et al., 2014)

Eptalofosuchus gen. nov.

Type species: Eptalofosuchus viridi sp. nov.

Etymology. Generic name from the Greek eptá for seven and lófos for hills, in allusion to Uberaba's nickname “cidade das sete colinas” (“the city of the seven hills”), plus souchus, Greek for crocodile, leading to Latin, suchus.

Eptalofosuchus viridi sp. nov.

Etymology. Specific name derives from the Latin viridi, meaning green, in allusion to the typical greenish color of the Uberaba Formation.

 Thiago S. Marinho, Agustín G. Martinelli, Giorgio Basilici, Marcus Vinícius T. Soares, André Marconato, Luiz C.B. Ribeiro and Fabiano V. Iori. 2021. First Upper Cretaceous notosuchians (Crocodyliformes) from the Uberaba Formation (Bauru Group), southeastern Brazil: Enhancing Crocodyliform Diversity. Cretaceous Research. In Press, 105000. DOI: 10.1016/j.cretres.2021.105000

  twitter.com/tupandactylus/status/1435982854891184133

 

[Ichthyology • 2022] Exon-capture Data and Locus Screening provide new insights into the Phylogeny of Flatfishes (Pleuronectoidei)

in Atta, Yuan, Li, ... et Tornabene, 2022.

  DOI: 10.1016/j.ympev.2021.107315

Highlights: 

• Phylogenetic analysis of flatfishes uses next-gen dataset with largest taxon sampling to date.

• Rhombosoleidae is found to be in Pleuronectoidea rather than Soleoidea.

• Positions of Poecilopsettidae and Scophthalmidae are dictated by tree-construction method.

• Many relationships in the flatfish tree are unstable due to periods of rapid evolution.

• 8/16 families need thorough analysis; 23/127 genera may not be monophyletic (21 not yet sampled).

Abstract

There is an extensive collection of literature on the taxonomy and phylogenetics of flatfishes (Pleuronectiformes) that extends over two centuries, but consensus on many of their evolutionary relationships remains elusive. Phylogenetic uncertainty stems from highly divergent results derived from morphological and genetic characters, and between various molecular datasets. Deciphering relationships is complicated by rapid diversification early in the Pleuronectiformes tree and an abundance of studies that incompletely and inconsistently sample taxa and genetic markers. We present phylogenies based on a genome-wide dataset (4,434 nuclear markers via exon-capture) and wide taxon sampling (86 species spanning 12 of 16 families) of the largest flatfish suborder (Pleuronectoidei). Nine different subsets of the data and two tree construction approaches (eighteen phylogenies in total) are remarkably consistent with other recent molecular phylogenies, and show strong support for the monophyly of all families included except Pleuronectidae. Analyses resolved a novel phylogenetic hypothesis for the family Rhombosoleidae as being within the Pleuronectoidea rather than the Soleoidea, and failed to support the subfamily Hippoglossinae as a monophyletic group. Our results were corroborated with evidence from previous phylogenetic studies to outline regions of persistent phylogenetic uncertainty and identify groups in need of further phylogenetic inference.

 

Keywords: Phylogenomics, Systematics, Soles, Flounders, Incomplete-lineage-sorting, Target-enrichment

Conclusions: 

Our analysis of flatfish systematics using an exon-capture dataset with relatively dense taxon sampling was mostly consistent with the leading phylogenetic hypotheses for the Pleuronectiformes (BHC model) and the Pleuronectidae (Vinnikov et al., 2018). We report a novel position for the family Rhombosoleidae and show extremely poor support for the subfamily Hippoglossinae. Collective inference using sixteen phylogenetic analyses demonstrates the tenuous nature of several flatfish relationships at various evolutionary scales, and how certain relationships are favored by particular methods. By examining our data within the historical context of flatfish systematics we were able to identify several regions where phylogenetic uncertainty is likely to remain and outline groups that should be targeted for further study.

 Calder J. Atta, Hao Yuan, Chenhong Li, Dahiana Arcila, Ricardo Betancur-R, Lily C. Hughes, Guillermo Ortí, Luke Tornabene. 2022. Exon-capture Data and Locus Screening provide new insights into the Phylogeny of Flatfishes (Pleuronectoidei). Molecular Phylogenetics and Evolution. 166,107315. DOI: 10.1016/j.ympev.2021.107315

 facebook.com/groups/204572449594711/posts/4564404300278149

[Herpetology • 2021] Duttaphrynus brevirostris, D. peninsularis & D. stomaticus • Lost, Forgotten, and Overlooked: Systematic Reassessment of Two lesser-known Toad Species (Anura, Bufonidae) from Peninsular India and Another Wide-ranging northern Species

A. Duttaphrynus brevirostris (BNHS 6126) from Kempholey Ghat region in Sakleshpur taluk. B. Duttaphrynus peninsularis (SDBDU 6370) from Wattakolli.
C–D. Duttaphrynus stomaticus: C. SDBDU 2015.2909 from Assam; D. SDBDU 2012.2170 from Rajasthan.

in Bisht, Garg, ... et , 2021. 

DOI: 10.3897/zse.97.61770

  facebook.com/BijuSDas

Abstract

We rediscovered two species of toads, Bufo stomaticus peninsularis and Bufo brevirostris, which were described from Peninsular India 84 and 101 years ago, respectively, but have not been reported since. Because the name-bearing types of both species are either damaged or lost, we provide detailed redescriptions, morphological comparisons, and insights into phylogenetic relationships with closely related members of the genus Duttaphrynus sensu lato, based on new material from the type locality of each species. We clarify and validate the identity of D. brevirostris, which was rediscovered from multiple localities in the Malenadu and adjoining coastal regions of Karnataka. We also demonstrate that Bufo stomaticus peninsularis, which was considered a synonym of Duttaphrynus scaber, is a distinct species. Bufo stomaticus peninsularis differs from Duttaphrynus scaber morphologically and genetically, and is more closely related to members of the Duttaphrynus stomaticus group. We also clarify the identity of the namesake species of the Duttaphrynus stomaticus group, which is reported widely in India and neighbouring countries, but lacks sufficient taxonomic information due to its brief original description and reportedly untraceable type material. We located and studied the complete syntype series of D. stomaticus, probably for the first time in over a century, and we report on the status of available specimens, provide detailed description of a potential type, compare it to related species, and clarify the species’ geographical range. Our molecular analyses suggest that D. stomaticus is minimally divergent from, and possibly conspecific with, D. olivaceus. Our analyses also clarify its relationship to the closely-related D. peninsularis comb. nov., with which it was previously confused. Finally, our study provides other insights into the phylogenetic relationships and genetic differentiation among various species of Duttaphrynus toads.

Key Words: Amphibia, Bufo stomaticus peninsularis, distribution, Duttaphrynus brevirostris, Duttaphrynus stomaticus group, Firouzophrynus, molecular phylogeny, redescription, rediscovery, taxonomy

Topotype of Duttaphrynus brevirostris (Rao, 1937), topotype of D. peninsularis (Rao, 1920), and referred specimens of D. stomaticus (Lütken, 1864) in life.
A. Duttaphrynus brevirostris (BNHS 6126) from Kempholey Ghat region in Sakleshpur taluk.
B. Duttaphrynus peninsularis (SDBDU 6370) from Wattakolli.
C–F. Duttaphrynus stomaticus: C. SDBDU 2015.2909 from Assam; D. SDBDU 2012.2170 from Rajasthan; E. SDBDU 2012.2172 from Delhi; and F. SDBDU 2012.2268 from Bihar.

Phylogenetic relationships and genetic differentiation in the genus Duttaphrynus.
 A. Maximum Likelihood phylogenetic tree based on 5,737 bp DNA comprising nine mitochondrial gene regions and two nuclear genes, showing phylogenetic relationships between the major species-level lineages. Values above and below the branches indicate Bayesian Posterior Probabilities (BPP) and RAxML Bootstrap Support (BS), respectively;
B. Maximum Likelihood barcoding tree based on 524 bp of the mitochondrial 16S rRNA sequences. BPP and BS support values are indicated above and below the branches, respectively. Coloured vertical bars outside the terminal node labels indicate putative species delimited in the bPTP analysis;
C. Median-Joining haplotype network based on 42 haplotypes recovered from 133 sequences of the 16S gene (420 bp). Size of the coloured circles is proportional to the number of haplotypes; black circles indicate median vectors; each branch represents a single mutation step; additional mutational steps are indicated by values in parentheses; photo credits: D. crocus (Guinevere O. U. Wogan), D. olivaceus (Parham Beyhaghi), and D. dhufarensis (Todd W. Pierson).

Duttaphrynus brevirostris (Rao, 1937)

Duttaphrynus peninsularis (Rao, 1920), comb. nov.

Duttaphrynus stomaticus (Lütken, 1864)

Geographical distribution of Duttaphrynus brevirostris (dark grey), D. peninsularis (blue), and D. stomaticus (orange).

Conclusions: 

The results of this study resolve long-standing uncertainty regarding the identities and taxonomic status of two toad species described from Peninsular India. Bufo brevirostris Rao, 1937 was considered a problematic taxon, because its original name-bearing types are lost. Bufo stomaticus peninsularis Rao, 1920 was long forgotten as an available name for Peninsular Indian populations closely related to Duttaphrynus stomaticus. We substantiate D. peninsularis to be a distinct species, which is both morphologically diagnosable and phylogenetically distinct. Taxonomic redefinition of both of these species was achieved not just by examining the original literature and available types, but also through an effort to rediscover new material from each species’ respective type locality. The redescription of Bufo brevirostris Rao, 1937 based on new topotypic material, along with detailed comparisons to related taxa, objectively clarifies its identification for future reference. Similarly, topotypic material for Bufo stomaticus peninsularis Rao, 1920 enabled a detailed re-evaluation of its taxonomic status in the absence of a well-preserved type. Altogether, our results emphasise that new collections from type localities of historically available names should be attempted when taxonomic resolution is not feasible on the basis of original descriptions or type specimens (Bailey 1933; Garg and Biju 2016).

The present work clarified the taxonomic identity of another species, Duttaphrynus stomaticus, which was overlooked due to its presumed wide distribution. This taxon was known only from its brief original description, and the available, original name-bearing types remained unexamined due to literature-based misconceptions concerning their untraceability (Dutta 1997; Ganesh et al. 2020). We located the well-preserved eight original type specimens, and clarified the status of name-bearing types and the identity of this species, which we redescribed to facilitate future taxonomic studies. This action also aided our objective of resolving the taxonomic status of D. peninsularis, which was originally defined as a variety of D. stomaticus. Our results have important implications concerning the taxonomy and geographical ranges of the two species. Hereafter, D. stomaticus should be considered as a species found in the northern regions of South Asia, whereas its sister taxon D. peninsularis should be recognised as a Peninsular Indian form (Fig. 4; Suppl. materal 1: Table S3). Detailed redescriptions provided in this study will enable proper identification and range delineation, and serve as the basis for future conservation action. Knowledge of phenotypic variation and phylogenetic affinities of both species will also facilitate a better understanding of patterns of genetic differentiation within the genus, particularly among the species of the Duttaphrynus stomaticus group.

   

 Karan Bisht, Sonali Garg, A. N. D. Akalabya Sarmah, Saibal Sengupta and S. D. Biju. 2021. Lost, Forgotten, and Overlooked: Systematic Reassessment of Two lesser-known Toad Species (Anura, Bufonidae) from Peninsular India and Another Wide-ranging northern Species. Zoosystematics and Evolution. 97(2): 451-470. DOI: 10.3897/zse.97.61770

  facebook.com/BijuSDas/posts/6017683671639383

  twitter.com/SDBiju1/status/1440376404089053207

[Herpetology • 2021] Cnemaspis balerion, C. lithophilis, C. regalis, C. smaug, C. wallaceii, etc. • Diversifying on the Ark: Multiple New Endemic Lineages of Dwarf Geckos (Squamata: Gekkonidae) from the Western Ghats provide insights into the Systematics and Biogeography of South Asian Cnemaspis

Coloration in life of all 12 holotypes; 
A: Cnemaspis balerion sp. nov.; B: C. lithophilis sp. nov.; C: C. rubraoculus sp. nov.; D: C. nimbus sp. nov.; E: C. wallaceii sp. nov.; F: C. smaug sp. nov.; 
G: C. regalis sp. nov.; H: C. galaxia sp. nov.; I: C. nigriventris sp. nov.;  J: C. flavigularis sp. nov.; K: C. palanica sp. nov.; L: C. jackieii sp. nov..

Pal, Mirza, Dsouza & Shanker, 2021

 DOI:  10.24272/j.issn.2095-8137.2021.074 

Photos by Saunak Pal.  facebook.com/SaunakPal 

Abstract

The highly speciose gekkonid genus Cnemaspis Strauch, 1887 is polyphyletic, with three distantly related and geographically isolated clades from Africa, South Asia (SA), and Southeast Asia. At present, there are 85 known species within SA Cnemaspis, although the number continues to increase rapidly with focused surveys and rigorous taxonomic work. Recent studies have provided valuable insights into the diversity and evolutionary history of SA Cnemaspis; however, most of these studies lack sufficient sampling in the Western Ghats (WG), where the genus has its greatest diversity. We addressed this research gap by conducting extensive sampling across the WG and re-examining museum specimens, thus providing a systematic account of various extant Cnemaspis species along with their distribution and natural history. We described 12 new species and a southern WG endemic clade of SA Cnemaspis. Ten of the newly described species are endemic to the forests of the southern WG. We also identified 10 well-supported subclades that can be separated across morphological, geographic, and phylogenetic axes. Time-calibrated phylogeny and ancestral area reconstructions confirmed the Paleocene origin of SA Cnemaspis in the WG and provide insights into its evolutionary history and biogeography. The discovery of multiple endemic and deeply divergent lineages further highlights the evolutionary significance of the WG for lizards.

Keywords: Biogeography,  Cnemaspis,  Dwarf geckos,  Molecular phylogeny,  New species,  Osteology,  South Asia,  Systematics,  Western Ghats

Coloration in life of all 12 holotypes
A: Cnemaspis balerion sp. nov.; B: Cnemaspis lithophilis sp. nov.; C: Cnemaspis rubraoculus sp. nov.; D: Cnemaspis nimbus sp. nov.; E: Cnemaspis wallaceii sp. nov.; F: Cnemaspis smaug sp. nov.; 
G: Cnemaspis regalis sp. nov.; H: Cnemaspis galaxia sp. nov.; I: Cnemaspis nigriventris sp. nov.;  J: Cnemaspis flavigularis sp. nov.; K: Cnemaspis palanica sp. nov.; L: Cnemaspis jackieii sp. nov..

Photos by Saunak Pal.

A: Cnemaspis balerion sp. nov.; B: Cnemaspis lithophilis sp. nov.; C: Cnemaspis rubraoculus sp. nov.; 
D: Cnemaspis nimbus sp. nov.; E: Cnemaspis wallaceii sp. nov.; F: Cnemaspis smaug sp. nov.

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I. wynadensis clade: 

Comprising nine known lineages viz. C. kolhapurensis; C. sisparensis (Theobald, 1876); C. kottiyoorensis Cyriac & Umesh, 2014; C. heteropholis Bauer, 2002; C. anaikattiensis Mukherjee, Bhupathy & Nixon 2005; C. wynadensis; C. chengodumalaensis Cyriac, Palot, Deuti & Umesh, 2020; C. zacharyi Cyriac, Palot, Deuti & Umesh, 2020; and C. magnifica Khandekar, Thackeray, Pal & Agarwal, 2020. We identified two new well-supported, distantly related lineages with moderate support, described here as Cnemaspis balerion sp. nov. and Cnemaspis lithophilis sp. nov. (Figure 3).

Cnemaspis balerion sp. nov.

Etymology: The species is named after “Balerion”, a famous dragon from George R. R. Martin’s epic fantasy novel series, ‘A song of Ice and fire’. The dragon was also known as ‘the black dread’ due to presence of black scales. Like Balerion, this new species shows the presence of distinct clusters of black scales forming spots on the dorsum. 

Suggested common name: Balerion forest gecko. 

Cnemaspis lithophilis sp. nov. 

Etymology: The species epithet is derived from the combination of the Greek word ‘lithos’ meaning stone and ‘philia’ meaning fondness, due to the close association of this species with rocks. 

Suggested common name: Sharavati rock gecko. 

I. wynadensis clade: A: Cnemaspis balerion sp. nov.;
 II. beddomei clade: C: C. rubraoculus sp. nov.; D: C. nimbus sp. nov.; E: C. wallaceii sp. nov.; F: C. smaug sp. nov.; G: C. regalis sp. nov.; H: C. galaxia sp. nov.; I: C. nigriventris sp. nov.;  

II. beddomei clade: 

Comprising six known lineages viz. C. beddomei; C. aaronbaueri Sayyed, Grismer, Campbell & Dileepkumar, 2019; C. anamudiensis Cyriac, Johny, Umesh & Palot, 2018; C. maculicollis Cyriac, Johny, Umesh & Palot, 2018; C. ornata (Beddome, 1870); and C. nairi Inger, Marx & Koshy, 1984. Seven of the newly identified lineages belong to this clade (Figure 2), described here as Cnemaspis rubraoculus sp. nov., Cnemaspis wallaceii sp. nov., Cnemaspis nimbus sp. nov., Cnemaspis smaug sp. nov., Cnemaspis nigriventris sp. nov., Cnemaspis galaxia sp. nov., and Cnemaspis regalis sp. nov. (Figure 3).

Cnemaspis rubraoculus sp. nov.

Etymology: The species epithet is derived by combining the Latin word ‘rubrum’, meaning red, and ‘oculus’ meaning eye; referring to the prominent red iris. 

Suggested common name: Ruby eyed forest gecko.

Cnemaspis nimbus sp. nov.

Etymology: The species epithet is derived from the Latin word ‘nimbus’ meaning cloud; referring to the prominent clouded pattern on the dorsum and flank. 

Suggested common name: Clouded forest gecko. 

Cnemaspis wallaceii sp. nov.

Etymology: The species epithet is a patronym, honouring Alfred Russel Wallace for his tremendous contribution to the field of biogeography. His work has been an inspiration for the authors and towards this study. 

Suggested common name: Wallace’s forest gecko.

Cnemaspis smaug sp. nov.

Etymology: The species is named after “Smaug”, the dragon from J. R. R. Tolkien's 1937 novel, ‘The Hobbit’. The name is derived from the old German verb ‘smeuganan’ meaning “to creep” or “to squeeze through a hole”. The type specimens of this species were found within crevices of rocks and boulders inside the forest. Additionally, like dragons, the dorsum is armoured with large conical tubercles. 

Suggested common name: Smaug forest gecko 

Cnemaspis regalis sp. nov.

Etymology: The species epithet is based on the Latin word ‘regalis’ meaning kingly or royal, due to the head ornamented with golden yellow crown like pattern in the males of this species. 

Suggested common name: Golden crowned day gecko or Royal day gecko.

Cnemaspis galaxia sp. nov.

Etymology: The species epithet is derived from the word ‘galaxy’ due to the dorsum colouration with the sun’s haze like yellow on the anterior and bluish white star like spots on a black background towards the posterior end, in the male of this species. 

Suggested common name: Galaxy day gecko. 

Cnemaspis nigriventris sp. nov.

Etymology: The species epithet is derived from the combination of Latin word ‘niger’ meaning black and ‘venter’ meaning belly, due to the distinct black ventral colouration in the male of this species. 

Suggested common name: Black-bellied day gecko.  

 

 

III. littoralis clade: 

Comprising two known lineages from the WG, including C. littoralis and the recently described C. palakkadensis Sayyed, Cyriac & Dileepkumar, 2020. Two new lineages belong to this clade, described here as Cnemaspis flavigularis sp. nov., and Cnemaspis palanica sp. nov. (Figure 3). This clade includes members of the C. podihuna clade from Sri Lanka, as observed in other recent studies (Agarwal et al., 2020b; Khandekar et al., 2019a).

Cnemaspis flavigularis sp. nov.

Etymology: The species epithet is derived from the combination of the Latin word ‘flavus’ meaning yellow and ‘gularis’ meaning throat, referring to the distinct yellow colouration of the throat in males of the new species. 

Suggested common name: Yellow throated day gecko. 

Cnemaspis palanica sp. nov.

Etymology: The species epithet refers to the ‘Palani Hills’ in the SWG of Tamil Nadu, from where this species is described. 

Suggested common name: Palani Hills day gecko.

 

G: Cnemaspis regalis sp. nov.; H: Cnemaspis galaxia sp. nov.; I: Cnemaspis nigriventris sp. nov.; 
 J: Cnemaspis flavigularis sp. nov.; K: Cnemaspis palanica sp. nov.; L: Cnemaspis jackieii sp. nov..

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IV. indica clade: 

Comprising three known lineages viz. C. indica; C. nilagirica Manamendra-Arachchi, Batuwita & Pethiyagoda, 2007; and C. anandani Murthy, Nitesh, Sengupta & Deepak, 2019.

V. bangara clade:

Comprising three known lineages viz. C. bangara; C. graniticola Agarwal, Thackeray, Pal & Khandekar, 2020; and C. yelagiriensis Agarwal, Thackeray, Pal & Khandekar, 2020.

VI. girii clade: 

Comprising seven known lineages viz. C. girii; C. mahabali Sayyed, Pyron & Dileepkumar, 2018; C. limayei Sayyed, Pyron & Dileepkumar, 2018; C. ajijae Sayyed, Pyron & Dileepkumar, 2018; C. flaviventralis Sayyed, Pyron & Dahanukar, 2016; C. amba Khandekar, Thackeray & Agarwal, 2019; and C. koynaensis Khandekar, Thackeray & Agarwal, 2019.

VII. goaensis clade: 

Comprising two known lineages viz. C. amboliensis Sayyed, Pyron & Dileepkumar, 2018 and C. goaensis.

VIII. mysoriensis clade: 

Comprising seven known lineages viz. C. mysoriensis (Jerdon, 1853); C. otai Das & Bauer, 2000; C. yercaudensis Das & Bauer, 2000; C. adii Srinivasulu, Kumar & Srinivasulu, 2015; C. avasabinae Agarwal, Bauer & Khandekar 2020; C. rishivalleyensis Agarwal, Thackeray & Khandekar 2020; and C. stellapulvis Khandekar, Thackeray & Agarwal, 2020.

IX. gracilis clade: 

Comprising four known lineages viz. C. gracilis; C. agarwali Khandekar, 2019; C. shevaroyensis Khandekar, Gaitonde & Agarwal, 2019; and C. thackerayi Khandekar, Gaitonde & Agarwal, 2019. A well-supported new lineage is described here as Cnemaspis jackieii sp. nov. (Figure 3).

Cnemaspis jackieii sp. nov.
E. female paratype [CESL 193], F. Habitat at type locality in Vairavankulam Reserve Forest, Tamil Nadu, G. communal egg deposition in used potter wasp nests.

Cnemaspis jackieii sp. nov.

Etymology: The species epithet is a patronym as a tribute to the famous actor and martial artist Jackie Chan. The new species was found to be very fast and moved rapidly on rock boulders, sneaking into the smallest crevices to escape when approached, reminiscent of the stunts of Jackie Chan. His many screen characters as an explorer and adventurer have been an inspiration for the authors. 

Suggested common name: Jackie’s day gecko. 

X. monticola clade: 

Comprising two known lineages viz. C. monticola and C. australis from the WG. This clade also includes members of the C. kandiana clade from Sri Lanka and Southeast Asian representatives of SA Cnemaspis, as observed in other recent studies (Agarwal et al., 2020b; Khandekar et al., 2019a).

   

CONCLUSIONS: 

Despite being one of the world’s “hottest biodiversity hotspots”, the WG have undergone substantial habitat loss and degradation due to changes in land-use patterns (Jha et al., 2000; Myers et al., 2000; Reddy et al., 2013). It has been suggested that significant parts of the WG will be urbanized by 2030 due to population expansion and development (Seto et al., 2012). The recent discoveries of new evolutionarily distinct lineages from the WG highlight the biogeographic and evolutionary significance of this region as well as the importance of sampling strategies to detect endemic lineages while conducting broad-level phylogenetic studies (Britz et al., 2020; Chaitanya et al., 2019; Dinesh et al., 2020; Mallik et al., 2019; Pal et al., 2018; Robin et al., 2017; Vijayakumar et al., 2019). The recognition of 10 new morphologically distinct and geographically isolated species of Cnemaspis from the SWG also highlights our lack of knowledge regarding the immense diversity of this region and its potential to harbor more unknown species. Recent discoveries of species that are morphologically similar to extant species within their respective clades (e.g., C. palakkadensis and C. littoralis; C. nilagirica and C. anandani, C. nairi and Cnemaspis nigriventris sp. nov.) further emphasize the potential for unexplored cryptic diversity in the WG. Most of the recently described evolutionarily distinct lineages, as well as those identified in this study, are range-restricted and have very specific microhabitat requirements. It is important to identify these unique habitats within the landscape and focus efforts on the conservation of unique lineages with diverse evolutionary histories.

In the last decade, there has been a surge in the discovery of new species of Cnemaspis from peninsular India, especially from regions not known to harbor high Cnemaspis diversity, such as the southern edge of the Mysore Plateau and the NWG. The molecular data generated in this study not only include new species, but also species previously known only from type localities or specimens, and thus provide a baseline for future taxonomic research. The discovery of a highly diverse SWG endemic clade (Clade II) and a dry zone granite-associated clade (Agarwal et al., 2020b) suggest that more effort is required to elucidate the diversity within SA Cnemaspis. Additionally, our findings stress the importance of spatial sampling and geographical distribution of species, along with molecular, phylogenetic, and morphological methods, in uncovering deeply divergent lineages.

Saunak Pal, Zeeshan A. Mirza, Princia Dsouza and Kartik Shanker. 2021. Diversifying on the Ark: Multiple New Endemic Lineages of Dwarf Geckos from the Western Ghats provide insights into the Systematics and Biogeography of South Asian Cnemaspis (Reptilia: Squamata). Zoological Research. 42(6); 675-691. DOI:  10.24272/j.issn.2095-8137.2021.074

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