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Friday, July 24th, 2020
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2:03a |
[Entomology • 2020] Veredatrypa gen. nov. • New Brazilian Tafaliscina increase the Diversity of this Neotropical Cricket Clade (Orthoptera: Gryllidae: Oecanthinae)
![](https://1.bp.blogspot.com/-2OChij4to70/Xxljb1FiSxI/AAAAAAAEGQw/H5bkd9dSH3gDw6bZDqX0kMUmYWLZUWyVACK4BGAYYCw/s400/Veredatrypa_rosai-novataxa_2020-Campos_Souza-Dias_et_Nihei--%2540Publi_MNHN.jpg) | Veredatrypa Campos Veredatrypa rosai Campos, Souza-Dias & Nihei, 2020.
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Tafaliscina Desutter, 1988 (Grylloidea Laicharting, 1781, Gryllidae Laicharting, 1781, Oecanthinae Blanchard, 1845, Paroecanthini Gorochov, 1986) are a Neotropical cricket clade with a remarkable morphological diversity. We study here their Brazilian representatives. We describe one new genus (Veredatrypa Campos n. gen.) and three new species from Cerrado and Caatinga, i.e., Veredatrypa rosai n. gen., n. sp., V. seca n. gen., n. sp. and V. fusca n. gen., n. sp., and two new species of Tafalisca Walker, 1869 from Amazon Forest, i.e., T. duckeana n. sp. and T. vestigialis n. sp. The new genus and the new species are mainly characterized by their male phallic complexes and forewings. An identification key of Tafaliscina Brazilian genera is provided, and the diversity of the subtribe is discussed in relation to the life habits and communication modalities of Tafaliscina genera.
KEYWORDS: Crickets, morphology, forewings, phallic complex, biodiversity, new genus, new species
![](https://3.bp.blogspot.com/-Lz1cxI3ok98/XxljWHn3v6I/AAAAAAAEGQo/yNBuyZBNj5IOZRtYeCksROkiywFj9nC-QCK4BGAYYCw/s400/Veredatrypa_rosai-novataxa_2020-Campos_Souza-Dias_et_Nihei--%2540Publi_MNHN.jpg) | Veredatrypa rosai n. gen., n. sp. adult female. |
SYSTEMATICS
Order OTHOPTERA Olivier, 1789 Superfamily Grylloidea Laicharting, 1781
Family Gryllidae Laicharting, 1781 Subfamily Oecanthinae Blancard, 1845
Tribe Paroecanthini Gorochov, 1986 Subtribe Tafaliscina Desutter, 1988
Veredatrypa Campos n. gen.
Type species. — Veredatrypa rosai n. gen, n. sp.
Species included. — Veredatrypa rosai n. gen., n. sp.; Veredatrypa seca n. gen., n. sp.; Veredatrypa fusca n. gen., n. sp.
Etymology. — Named after “Grande Sertão: Veredas”, a novel written by the Brazilian novelist João Guimarães Rosa. Grande Sertão: Veredas (in English, translated as The Devil to Pay in the Backlands) is one of the masterpieces of Brazilian literature and one of the most important novels in Portuguese language literature due to its style and complexity. Grande Sertão: Veredas is also a Brazilian National Park that protects a large fragment of Brazilian Cerrado, including wetlands areas known as “veredas”. That Park is located in the same area where the novel happened and is the type-locality of this new genus.
Distribution. — This genus is recorded for Brazilian open forests as the Cerrado and Caatinga, in States of Minas Gerais, Goiás and Ceará.
Diagnosis. — The genus is separated from the other genera of Tafaliscina by the following characters: median ocelli very reduced, almost no discernible; DD with longitudinal lateral bands from cephalic to caudal margin. Male: metanotum with two pairs of projections, with cluster of bristles (absent in V. seca n. gen., n. sp.). FWs with anal vein area slightly bulged dorsally, stridulatory file surrounded by strong sclerotization, hv anterior region strongly sclerotized. Male genitalia: MLophi triangular, with a pointed apex; LLophi very short or absent; EctF strongly sclerotized, wide anteriorly, thin posteriorly, going along MLophi, apex folding dorsally, connecting to the apex of MLophi; endophallic apodeme bifid. Female: ovipositor short than cerci, with two lateral protuberances in dorsal and ventral views. Female genitalia: copulatory papilla distal half constricted, without furrow ventrally.
Veredatrypa rosai n. gen., n. sp.
Etymology. — Name after the Brazilian novelist João Guimarães Rosa, author of the novel “Grande Sertão: Veredas”.
Veredatrypa seca n. gen., n. sp.
Etymology. — The word “seca” means “dry” in Portuguese, referring to the climatic conditions of the region where this species was collected. The name is a noun in apposition.
Veredatrypa fusca n. gen., n. sp.
Etymology. — From Latin, “fusca” means “dark”. This species is darker than the other species of the genus.
Tafalisca duckeana n. sp.
Etymology. — Referring to “Reserva Florestal Adolpho Ducke”, a forest area managed by the Instituto Nacional de Pesquisas da Amazônia (INPA), where this species was discovered.
Tafalisca vestigialis n. sp.
Etymology.— From Latin, “vestigialis” means “vestigial”, referring to the vestigial stridulatory file of this species.
Lucas Denadai de Campos, Pedro G. B. Souza-Dias and Silvio Shigueo Nihei. 2020. New Brazilian Tafaliscina increase the Diversity of this Neotropical Cricket Clade (Orthoptera: Grylloidea: Gryllidae: Oecanthinae: Paroecanthini). ZOOSYSTEMA. 42(19); 331-353. DOI: 10.5252/zoosystema2020v42a19
| 4:11a |
[PaleoMammalogy • 2020] Eomakhaira molossus • A New Saber-Toothed Sparassodont (Metatheria: Thylacosmilinae) from the Early Oligocene (?Tinguirirican) Cachapoal Locality, Andean Main Range, Chile ![](https://1.bp.blogspot.com/-WhNDwNSyR8A/Xxle1WIsChI/AAAAAAAEGQc/IeSPpx-1VmomjCVinOLnSJym6R1hCQsBgCK4BGAYYCw/s400/Eomakhaira_molossus-novataxa_2020-Engelman_Flynn_Wyss_et_Croft__%2540DCPaleo.jpg) | Eomakhaira molossus
Engelman, Flynn, Wyss & Croft, 2020
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Abstract Thylacosmiline sparassodonts (previously recognized as thylacosmilids) are among the most iconic groups of endemic South American Cenozoic mammals due to their distinctive morphology and convergent resemblance to saber-toothed placental carnivores. However, the early evolution of this group and its relationship to other sparassodonts remains poorly understood, primarily because only highly specialized Neogene taxa such as Thylacosmilus, Anachlysictis, and Patagosmilus are well known. Here, we describe a new Paleogene sparassodont, Eomakhaira molossus, from the Cachapoal locality of central Chile, the first sparassodont reported from early Oligocene strata of the Abanico Formation. Eomakhaira shares features with both Neogene thylacosmilines and Paleogene “proborhyaenids,” and phylogenetic analyses recover this taxon as sister to the clade of Patagosmilus + Thylacosmilus. This broader clade, in turn, is nested within the group conventionally termed Proborhyaenidae. Our analyses support prior hypotheses of a close relationship between thylacosmilines and traditionally recognized proborhyaenids and provide the strongest evidence to date that thylacosmilines are proborhyaenids (i.e, the latter name as conventionally used refers to a paraphyletic group). To reflect the internestedness of these taxa, we propose use of Riggs' (1933) original name Thylacosmilinae for the less inclusive grouping and Proborhyaenidae for the more inclusive one. Saber teeth arose just once among metatherians (among thylacosmilines), perhaps reflecting a developmental constraint related to nonreplacement of canines in metatherians; hypselodonty may have relaxed this potential constraint in thylacosmilines. The occurrence of Eomakhaira in strata of early Oligocene age from the Chilean Andes demonstrates that the stratigraphic range of thylacosmilines spanned almost 30 million years, far surpassing those of saber-toothed placental lineages.
![](https://4.bp.blogspot.com/-79L0jasO9lc/Xxlc25bZBuI/AAAAAAAEGP4/4RNApSV4AkstkfhyfOFNWHUE0a6d0eOlQCK4BGAYYCw/s400/Eomakhaira_molossus-novataxa_2020-Engelman_Flynn_Wyss_et_Croft__%2540DCPaleo.jpg) | FIG 3. Photographs and CT segmentation of the holotype of Eomakhaira molossus, a partial skull of a senescent individual preserving the rostrum and the anterior portion of the mandible (SGOPV 3490) in left (A, B) lateral views. In renderings of the CT segmentation, nasal in orange, facial process of the lacrimal in teal, palatine in blue, all other bones of the cranium (maxilla, jugal, frontal, etc.) in purple, teeth in yellow, and dentary in green. Scale = 30 mm. |
SYSTEMATIC PALEONTOLOGY
MAMMALIA Linnaeus, 1754 METATHERIA Huxley, 1880 SPARASSODONTA Ameghino, 1894 BORHYAENOIDEA Simpson, 1930 PROBORHYAENIDAE Ameghino, 1897
Phylogenetic Definition: Proborhyaenidae refers to all sparassodonts more closely related to Proborhyaena gigantea than to Borhyaena tuberata, Prothylacynus patagonicus, Lycopsis torresi, Cladosictis patagonica, or Sipalocyon gracilis. This is a stem-based definition (de Queiroz and Gauthier, 1990).
THYLACOSMILINAE Riggs, 1933
Phylogenetic Definition: Thylacosmilinae refers to all sparassodonts more closely related to Thylacosmilus atrox than to Proborhyaena gigantea, Borhyaena tuberata, Prothylacynus patagonicus, Lycopsis torresi, Cladosictis patagonica, or Sipalocyon gracilis. This is a stembased definition (de Queiroz and Gauthier, 1990).
Eomakhaira molossus, gen. et sp. nov.
Etymology: The name of the genus derives from the Greek root Eos, meaning “dawn,” and makhaira, a type of short sword or large knife (often translated as “carving knife”), in reference to the bladelike canines of thylacosmilines. The specific epithet comes from the Greek molossus, a term used to refer to short-snouted, robust-skulled dog breeds such as mastiffs and bulldogs and refers to the short, robust snout of this species. Gender is masculine.
Diagnosis: A member of Borhyaenoidea based on its short, robust rostrum, presence of lingual median canine sulci, extremely small protocone, small and unicuspid talonid on m4. Differs from all other borhyaenoid sparassodonts in the following combination of features: small size (smaller than most other borhyaenoids; length of m1–4 = 37.3 mm, comparable to Fredszalaya hunteri or the extant dasyuromorphian Sarcophilus harrisii); maxilla very deep and maxillary “cheeks” absent; mandibular symphysis unfused and anteroposteriorly narrow; two mental foramina present; length/width ratio of palate >1.5; palate extending to level of M4; presence of postpalatine tori (shared only with Arminiheringia and possibly Callistoe among borhyaenoids); absence of postpalatine torus foramen; sphenorbital foramen opening dorsal to M4; large canines; absence of longitudinal striations on the canine roots (shared only with other thylacosmilines and possibly Lycopsis viverensis); median keel on the labial face of upper canines; medial sulcus on lingual face of upper and lower canines; short lower canine roots; presence of three premolars with no diastemata between them; premolars large and robust but not globular; asymmetric protoconid of P1 (shared only with Arminiheringia and Callistoe); P3 significantly longer than p3 (possibly autapomorphic for this taxon); bulbous roots only on p3; preparacingulum absent; M3 with narrow stylar shelf and prominent ectoflexus; M4 extremely narrow anteroposteriorly (only comparable to Patagosmilus among borhyaenoids), subequal or greater in width to M3, and with three roots; protocone vestigial (at least on M4); absence of an anteriorly projecting ventral keel of paraconid (which only occurs in proborhyaenids among sparassodonts); protoconid of m4 posteriorly salient; metaconid absent on m4 and probably m2–3; posterolabial cingulid present; talonid of m4 almost absent; and p1–3 short relative to m1–4 (shared with Paraborhyaena among borhyaenoids with three premolars). Canines more mediolaterally compressed than in borhyaenoids other than Patagosmilus, Thylacosmilus, and possibly Proborhyaena. P/p3 labiolingually narrower than in Fredszalaya, Plesiofelis, Acrocyon, Arctodictis, Australohyaena, Borhyaena, and Callistoe, but wider than in Prothylacynus and some individuals of Pharsophorus, comparable in relative proportions to Arminiheringia, Paraborhyaena, and Proborhyaena.
![](https://2.bp.blogspot.com/-hMjHL1bCBJw/Xxldcc0NPOI/AAAAAAAEGQQ/YyWCDnFmp2wlVo473T63QxvfCcurKnbuQCK4BGAYYCw/s400/Eomakhaira_molossus-novataxa_2020-Engelman_Flynn_Wyss_et_Croft__%2540DCPaleo.jpg) | FIG. 19. Size comparison of representative Paleogene proborhyaenids. From largest to smallest, Proborhyaena gigantea (in blue), the largest known proborhyaenid (scaled after AMNH 29576, the largest specimen of this taxon); Callistoe vincei (in green), the smallest named proborhyaenid prior to this study (scaled after the holotype specimen, PVL 4187); Eomakhaira molossus (in red), scaled after SGOPV 3490. Homo sapiens (170 cm tall) to right for comparison. Body mass for Proborhyaena from Prevosti et al. (2013) and Croft et al. (2018), Callistoe from Argot and Babot (2011), and Eomakhaira from the present study. Silhouettes for Proborhyaena (CC-BY-SA 3.0), duplicated for Callistoe and Eomakhaira, and Homo (CC0 1.0) by Zimices, and NASA, respectively, from PhyloPic. Scale bar = 1 m. |
![](https://3.bp.blogspot.com/-ZAGG0fjSHNA/XxldA1p8MYI/AAAAAAAEGQA/Y69ReFonWrYbKWTyFwgg71j3YJNGCtmAACK4BGAYYCw/s400/Eomakhaira_molossus-novataxa_2020-Engelman_Flynn_Wyss_et_Croft__%2540DCPaleo.jpg) | FIG. 21. Temporal durations of major lineages of mammalian saber-toothed carnivores, with metatherian lineage in grey and placental lineages in black. A representative skull of each clade is depicted to the right; from top to bottom: Thylacosmilus atrox (Thylacosmilinae), Machaeroides eothen (Machaeroidinae), Hoplophoneus primaevus (Nimravidae), Barbourofelis fricki (Barbourofelidae) and Smilodon fatalis (Machairodontinae). Images of Machaeroides, Barbourofelis, and Smilodon modified from Antón (2013), Thylacosmilus from Riggs (1934), and Hoplophoneus from Scott and Jepsen (1936) and Bryant (1996). Tick mark on Thylacosmilinae record represents the oldest occurrence of this clade (~20.2 Ma) prior to the discovery of Eomakhaira. Abbreviations: Plio., Pliocene; Ple., Pleistocene. |
Russell K. Engelman, John J. Flynn, André R. Wyss and Darin A. Croft. 2020. Eomakhaira molossus, A New Saber-Toothed Sparassodont (Metatheria: Thylacosmilinae) from the Early Oligocene (?Tinguirirican) Cachapoal Locality, Andean Main Range, Chile. American Museum Novitates. 2020(3957); 1-75. DOI: 10.1206/3957.1
| 4:50a |
[Botany • 2020] Begonia cabanillasii (section Baryandra, Begoniaceae) • A New Species from El Nido, Palawan, the Philippines
![](https://2.bp.blogspot.com/-8cUPXIgxhJo/XxlsdQiFadI/AAAAAAAEGRg/5fRuDW-fDNINtBaUjT6bgQhJ-NgCdXqsQCK4BGAYYCw/s400/Begonia_cabanillasii-novataax_2020-Ang_Tandang_Agcaoilli_et_Bustam__%2540PhTaxa.jpg) | Begonia cabanillasii Y.P.Ang, Tandang, J. Agcaoilli et R. Bustam.
in Ang, Tandang, ... et Bustamante, 2020. |
Abstract While assisting El Nido-Taytay Managed Resource Protected Area in their biodiversity monitoring of El Nido, Palawan, the authors came across a very distinctive and small population of Begonia. Based on careful investigation, the unknown species is confirmed as new to science, and is the latest addition to the species rich section Baryandra. A new species, Begonia cabanillasii is hereby described and illustrated. It is compared with phenetically similar species B. suborbiculata. Based on guidelines by IUCN, the new species is proposed to be Critically Endangered (CR) C2a.
Keywords: Eudicots, Begonia suborbiculata, Begonia quinquealata, Malesia, plant taxonomy
Yu Pin Ang, Danilo N. Tandang, John Michael M. Agcaoili and Rene Alfred Anton Bustamante. 2020. Begonia cabanillasii (section Baryandra, Begoniaceae), A New Species from El Nido, Palawan, the Philippines. Phytotaxa. 453(3); 244–254. DOI: 10.11646/phytotaxa.453.3.6 | 10:19a |
[Invertebrate • 2020] Borgesminthurinus andinus • A New Genus and Species of Katiannidae (Collembola: Symphypleona) from Bolivia
![](https://2.bp.blogspot.com/-vRCeoGItIf0/Xxq0oioXt7I/AAAAAAAEGU4/YWOlncKbs-Qx1t7uhIUXk_a7jqCgUQ4FACK4BGAYYCw/s400/Borgesminthurinus_andinus-novataxa_2020-Palacios-Vargas_et_Vacaflores-Argando%25C3%25B1a__zootaxa_4819-2.jpg) | Borgesminthurinus andinus
Palacios-Vargas & Vacaflores-Argandoña, 2020.
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Abstract The new genus Borgesminthurinus gen. nov. from Bolivia shares with Sminthurinus the presence of antennal segment IV undivided, antennal segment III with one papilla; sacs of ventral tube smooth; each tenacular rami with 3 teeth and a basal appendix. They also have dens with ventral chaetotaxy reduced and lack mucronal seta; but new genus clearly differs in having thick and barbulate setae on head and body, lacking neosminthuroid setae on abdomen and the presence of seta a0 on Abd. VI acuminate. The new genus differs from Katianna which has divided antennal segment IV, vertex of head with spine-like setae and setae of the body long and smooth.
Keywords: Systematics, Neotropical species, La Paz, Collembola
![](https://1.bp.blogspot.com/-UKgG-PLvueg/Xxqy3gyustI/AAAAAAAEGUg/jB1PB_ZyzYsPaOliefBeCPEOpfcYyzqcQCK4BGAYYCw/s400/Borgesminthurinus_andinus-novataxa_2020-Palacios-Vargas_et_Vacaflores-Argando%25C3%25B1a__zootaxa_4819-2.jpg) | Borgesminthurinus andinus sp. nov.: habitus. |
Systematics Order Symphyleona Börner, 1901 Superfamily Katiannoidea Bretfeld, 1994 Family Katiannidae Börner, 1913
Borgesminthurinus gen. nov. Palacios-Vargas
Diagnosis. Body globular, characterized by having thick, relatively long and slightly barbulate setae, very abundant on head and body; with full set of trichobothria present; A, B, C in a triangle on great abdomen and one pair of trichobothria on lesser abdomen (D); no neosminthuroid setae. Setae in large abdomen are thick but short. in singlets. Normal chewing mouth parts. Ant. IV undivided and longer than Ant. III; Ant. III with papilla divided in lobes. Tibiotarsi with 7 tenent hairs on distal whorl; unguis with pseudonychia and no tunica. Sacs of ventral tube smooth; tenacular rami with 3 teeth and basal appendix, corpus tenaculum with 2 setae. Metathoracic legs with trochanteral organ. Dens with ventral chaetotaxy reduced (formula 4, 2 … 1); mucro small, without seta, internal edge crenulate, external smooth. Seta a0 of Abdominal segment VI of females no furcate and no winged setae on lateral anal valves; subanal appendix of female palmate.
Type species. Borgesminthururinus andinus gen. nov. sp. nov.
Etymology. The new genus is dedicated to Jorge Luis Borges (Buenos Aires, August 24, 1899‒‒Geneva, Switzerland, June 14, 1986), the Argentinean writer, author of the Manual of Fantastic Zoology, which included spherical animals as Symphypleona are
Borgesminthurinus andinus gen. nov., sp. nov. Palacios-Vargas et Vacaflores-Argandoña
Derivatio nominis: The name of the new species is locative, for Los Andes.
José G. Palacios-Vargas and Ma. René Vacaflores-Argandoña. 2020. A New Genus and Species of Katiannidae (Collembola: Symphypleona) from Bolivia. Zootaxa. 4819(2); 201–215. DOI: 10.11646/zootaxa.4819.2.1
| 11:09a |
[Entomology • 2020] Taxonomic Study of Lebinthus Stål, 1877 (Orthoptera: Gryllidae: Eneopterinae) with Description of Six New Species in the Philippines
![](https://4.bp.blogspot.com/-bW4VKxy-3CA/Xxq-NlR1mCI/AAAAAAAEGWs/kxK23Ewo5Vs2v8jmue7V23QjY75D1YZggCK4BGAYYCw/s400/Lebinthus_bitaeniatus-St%25C3%25A5l__2020-Baroga-Barbecho_Tan_et-al--%2540ISYEBsp.png) | Lebinthus bitaeniatus Stål, 1877
in Baroga-Barbecho, Tan, et al., 2020. |
Abstract The genus Lebinthus Stål, 1877 (Orthoptera: Gryllidae: Eneopterinae: Lebinthini) in the Philippines is studied. New locality records are provided for L. bitaeniatus Stål, L. sanchezi Bolívar, L. polillensis Baroga, Yap & Robillard, L. puyos Robillard, and L. luae Robillard & Tan. Six new species are described from Mt. Porras, Sibalom (L. dannybaletei sp. nov.), Mt. Balatukan, Misamis Oriental (L. parvus sp. nov.), Mt. Empagatao, Misamis Oriental (L. palaceus sp. nov.), Tarumpitao, Palawan (L. hamus sp. nov.), Mt. Malinao, Albay (i sp. nov.), and Boracay, Aklan (L. boracay sp. nov.). A distribution map, a taxonomic key, and an updated checklist of the genus in the Philippines are also provided.
Keywords: Orthoptera, checklist, distribution, key, Lebinthini, redescription, Southeast Asia, taxonomy
![](https://4.bp.blogspot.com/-gABbMGBPNpc/Xxq97i6d8CI/AAAAAAAEGWg/CDN2DeCEqk0HSd_BPy0jTeVr8amVjMPsQCK4BGAYYCw/s400/Lebinthus_sanchezi-Bol%25C3%25ADvar__2020-Baroga-Barbecho_Tan_et-al--zootaxa_4816-4.jpg) | Lebinthus sanchezi male adult in its natural environment in Paete (Luzon). |
![](https://4.bp.blogspot.com/-bW4VKxy-3CA/Xxq-NlR1mCI/AAAAAAAEGWs/kxK23Ewo5Vs2v8jmue7V23QjY75D1YZggCK4BGAYYCw/s400/Lebinthus_bitaeniatus-St%25C3%25A5l__2020-Baroga-Barbecho_Tan_et-al--%2540ISYEBsp.png) | Lebinthus bitaeniatus male and female adults in pre-copulatory mode in their natural environment in Mount Makiling (Luzon). |
Lebinthus dannybaletei Baroga-Barbecho & Robillard sp. nov.
Etymology. The name of the species is dedicated to the late Mr. Danilo “Danny” S. Balete, a well-known Filipino Mammalogist and conservationist, for his contribution to science and Philippine conservation. He was the mentor of JBB in the mentoring program of the Biodiversity Conservation Society of the Philippines during its Annual Scientific Conference in 2010.
Lebinthus parvus Baroga-Barbecho & Robillard sp. nov.
Etymology. The name comes from the Latin word “parvus” which means “small” referring to the size of the species, which is the smallest Lebinthus among Mindanao species.
Lebinthus palaceus Baroga-Barbecho & Robillard sp. nov.
Etymology. The name palaceus is an adjective which means “spade-shaped”, referring to the shape of the pseudepiphallic paramere of the male genitalia.
Lebinthus hamus Baroga-Barbecho & Robillard sp. nov.
Etymology. The name comes from a Latin word “hamo” which means “hook” referring to the shape of the pseudepiphallic parameres and sclerites in ectophallic arc of male genitalia.
Lebinthus magayon Baroga-Barbecho & Robillard sp. nov.
Etymology. The name comes from the word “magayon” of the Bicol language, which is a term used to appreciate the beauty of a woman, relating it to the type material used to describe the species.
Lebinthus boracay Baroga-Barbecho, Yap & Robillard sp. nov.
Etymology. The species is named after the type locality.
Jessica B. Baroga-Barbecho, Ming Kai Tan, Sheryl A. Yap and Tony Robillard. 2020. Taxonomic Study of Lebinthus Stål, 1877 (Orthoptera: Gryllidae: Eneopterinae) with Description of Six New Species in the Philippines. Zootaxa. 4816(4); 401–438. DOI: 10.11646/zootaxa.4816.4.1
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