Research Article Discovering insect species based on photographs only: Journal of Orthoptera Research 2021, 30(2): 173-184 The case of a nameless species of the genus Scaria (Orthoptera: Tetrigidae) Niko KaAsALo!, Maks DerRANJA!2, KARMELA Apzic!2, RoBERTO SINDACC*, Josip SkEJo!23 1 Department of Biology, Division of Zoology, Evolution Lab, University of Zagreb, Faculty of Science, Rooseveltov trg 6, HR-10000 Zagreb, Croatia. 2 SIGTET—Special Interest Group Tetrigidae, Zagreb, Croatia. 3 IUCN/SSC Grasshopper Specialist Group, Zagreb, Croatia. 4 c/o Museo Civico di Storia naturale, via San Francesco di sales 88, I-10022 Carmagnola, Torino, Italy. Corresponding authors: Niko Kasalo (niko.kasalo5@gmail.com), Josip Skejo (jskejo@biol.pmf.hr) Academic editor: Maria-Marta Cigliano | Received 12 March 2021 | Accepted 27 July 2021 | Published 14 December 2021 http://zoobank.org/98819DBB-E028-4BA0-A623-A4AF0270CEF4 Citation: Kasalo N, Deranja M, Adzic¢ K, Sindaco R, Skejo J (2021) Discovering insect species based on photographs only: The case of a nameless species of the genus Scaria (Orthoptera: Tetrigidae). Journal of Orthoptera Research 30(2): 173-184. https://doi.org/10.3897/jor.30.65885 Abstract A heated debate on whether a new species should be described without a physical specimen, i.e., by designating a photographed specimen to serve as a holotype, has been ongoing for a long time. Herewith, without no- menclatural actions, a new species of the Batrachidein pygmy grasshoppers belonging to the genus Scaria Bolivar, 1887 is identified from the Andean rainforest in Peru. This species is clearly different from all its congeners by morphology and coloration. Two individuals of this peculiar species are known only from the photographs found on iNaturalist. The species has not been observed since 2008 when the photographs were taken. A short historical overview of the topic is given, illustrating the pros and cons of photograph-based species description. The concepts of names, holotypes, research effort, and conservation are discussed and related to the problem at hand. The current state of the taxonomic community’s beliefs regarding this issue is reflected by the authors’ three unsuccessful attempts to name this new species. Keywords Amazon, conservation, new species, Orthoptera, Peruvian Yungas, photog- raphy-based taxonomy, pygmy grasshopper, the ICZN Introduction Describing species from photographs or illustrations is not a common practice, but it has been done a number of times (Fricke and Kacher 1982, Welch et al. 1986, Jonkers and Roersma 1990, Wallach and Jones 1992, Jones et al. 2005, Sinha et al. 2005, Robb et al. 2013, Cheng et al. 2015, Marshall and Evenhuis 2015, In- grisch et al. 2016, Lonsdale and Marshall 2016, Nardelli 2016). Most of the names based on photographs are valid, but excep- tions do exist (for an extensive historical overview of this practice, see Krell and Marshall (2017)). Some of these publications have sparked a debate about the validity of describing species in this way. The following paragraphs provide a brief overview of the cur- rent state of the debate. Some authors (Timm and Ramey 2005, Santos et al. 2016) have implied that the International Code of Zo- ological Nomenclature (ICZN 1999) prohibits naming a species without a physical holotype, which is not correct (see Discussion). While Marshall and Evenhuis (2015) agree that collecting specimens is “highly desirable” as it allows for consideration of their internal morphology, microscopic and genetic characters and preservation of data for future access, they argue that, in certain cases, the specimens cannot be preserved and are getting harder to come by due to rigorous restrictions on collecting and trans- porting biological material, finally concluding that “collecting specimens is highly desirable, but it is indeed no longer required” because there is a growing number of high-quality photographs of taxa made by “digital collectors.” A prediction is made that “this situation will inevitably force the biodiversity community to adapt to growing numbers of new taxa recognized without ben- efit of dead, preserved type specimens” (Marshall and Evenhuis 2015). L6bl et al. (2016) consider such a practice dangerous, as it promotes quick capturing of visual data instead of careful observa- tion. They fear that works describing taxa based on an incomplete set of characters (i.e., photographs) may encourage non-experts to try to publish their works, harming the whole field of taxonomy as a result. These fears are considered unfounded by Shatalkin and Galinskaya (2017), as there are indeed very rigid requirements (ICZN 1999) for describing species from photographic evidence. Marshall and Evenhuis (2015) cite Minteer et al. (2014) in claim- ing that “collecting specimens is no longer required to describe a species...” The work in question, though, presents a limited set of examples (birds, amphibians, and plants) where scientists played a role in the extinction of those animals and suggests using alter- native means of sampling endangered species through high-res- olution photography, audio recording, and nonlethal sampling. The case has never been made for insects, and Marshall and Even- huis (2015) themselves agree that such a case would be difficult to make. Whether the cases illustrated by Minteer et al. (2014) even represent the issue was called into question by Krell and Wheeler (2014), but this is beyond the scope of our paper. L6bl et JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) 174 al. (2016) suggested publishing information about the existence of a new species without naming it. This is certainly possible but effectively discouraged due to the difficulty of finding a journal that would publish such a finding (Amorim et al. 2016). Having exhausted all the alternatives, we do exactly that. Amorim et al. (2016) also suggest that, while being a fine addition to the process of describing a species, photographic evidence cannot replace the usual process of “collecting, prepar- ing, comparing, describing, and delimiting species, that allows identifications (which are always hypotheses) to be double- checked.” We agree with this view but also feel the need to claim that there are and should be certain exceptions where blindly following this protocol could rob the scientific community of valuable knowledge. The immense administrative complexity of establishing a new species based on photographs has prevented us from doing so. An example to illustrate how photography- based taxonomy may introduce chaos is Presbytis johnaspinalli Nardelli, 2015, a new monkey species in the subfamily Colobi- nae, described from pictures of caged animals found on the In- ternet (Nardelli 2015). The morphological characters provided were not sufficient to clearly differentiate it from similar species, and some (Nijman 2015) have even suggested that the mon- keys were bleached by the traders who were trying to make them more visually appealing. Nardelli (2016) responded to the criti- cism, defending his claims, but the status of Presbytis johnaspi- nalli remains unclear. In the case of Scaria sp., it is clear that this is a hitherto unknown species. Record of a new unnamed Scaria N. KASALO, M. DERANJA, K. ADZIG, R. SINDACO AND J. SKEJO In this paper, we describe an unnamed species of a pygmy grasshopper (family Tetrigidae, subfamily Batrachideinae Boli- var, 1887), belonging to the tribe Batrachideini and genus Scaria Bolivar, 1887 (according to Cigliano et al. 2020) based on two photographs posted on the iNaturalist website by one of the authors (Roberto Sindaco; https://www.inaturalist.org/observa- tions/9968031) and three previously unpublished photographs taken by Roberto Sindaco at the same time as the original two. The photographs, taken in August 2008 in Peru (Fig. 1), were posted in 2018, and the specimens photographed were identified by the sen- ior author as a new species belonging to the subfamily Batrachid- einae. Since the photographs were posted, no one has described the species, nor has there been another recorded sighting of it. Differences in morphology observed from the five photographs may be sufficient to propose a new species within the genus Scaria Bolivar, 1887. Despite that, we decide not to establish one. The taxonomic community does not accept photography-based taxon- omy for reasons developed above. This is proven by rejections of this very manuscript when the naming of the species was includ- ed. Thus, this study aims to report an interesting unknown Scaria without nomenclatural actions. The specimens of the unnamed species were found in Peru (Bongara Province, Department of Am- azonas, Peru), which is a part of the Neotropical biogeographic region. This area is home to an astonishing number of animal and plant species (Elton 1973), owing such diversity to favorable envi- ronmental factors and long evolutionary history (Condon 2008). Considering these facts, it is clear that there is a wealth of species Tropical, rainforest (Af) Tropical, monsoon (Am) Tropical, savannah (Aw) Arid, desert, hot (BWh) Arid, desert, cold (BWk) Arid, steppe, hot (BSh) Arid, steppe, cold (BSk) Temperate, dry summer, warm summer (Csb) Temperate, dry summer, cold summer (Csc) Temperate, dry winter, warm summer (Cwb) Temperate, dry winter, cold summer (Cwc) Temperate, no dry season, warm summer (Cfb) Temperate, no dry season, cold summer (Cfc) Polar, tundra (ET) Polar, frost (EF) Fig. 1. Position of the only known locality of Scaria sp. Peroles near Yambrasbamba, marked with a star on the map of Peru with an- notated K6ppen-Geiger climate classification. Adapted from Beck et al. (2018). JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) N. KASALO, M. DERANJA, K. ADZIC, R. SINDACO AND J. SKEJO unknown to science that are waiting to be described. Unfortunate- ly, this impossible task (due to its sheer volume) is becoming even more discouraging due to the ongoing destruction of the entire region (Kehoe 2019). Drastic measures should be taken to reduce our negative impact on the environment because stable ecology is not only the playground of taxonomists, but also the corner- stone of the sustainability of life itself (Brown 1997, Kehoe 2019). The problem of declining biodiversity (Leather 2017, Forister et al. 2019) is only growing more significant, and cases such as this will keep happening. The unwillingness of the taxonomic community to discuss this issue and arrive at a set of rules that will apply to everybody equally is a major obstacle for scientists who find them- selves in possession of publishable material in this field. With this paper, we hope to take a step towards an eventual solution. Materials and methods Locality information.—The unnamed Scaria species is, for now, known only from a single locality in Peru. Peroles, near Yambras- bamba and 1905 m above sea level [5.670300°S, 77.918900°W], is one of the northernmost rainforests of the Peruvian Yungas and connects Amazon and Andes, a peculiar region where mosaics of unique rainforests reach extremely high altitudes. This ecoregion is considered to be in an almost critically endangered state (Beck 2018, WWE 2020). Although the locality is very close to two pro- tected areas—the Cordillera Colan Natural Sanctuary and the Alto Mayo protected forest—the area is severely deforested and looks like a mosaic of patches of forest alternating with deforested areas for grazing cattle. This forest is known for hosting the critically en- dangered yellow-tailed woolly monkey (Oreonax flavicauda (Hum- boldt, 1812)), one of “The World’s 25 Most Endangered Primates” (Shanee et al. 2019). Taxonomy.—Taxonomy follows Orthoptera Species File (OSF; Cigliano et al. 2020), while nomenclature is in accordance with the International Code of the Zoological Nomenclature (ICZN 1999). Systematics of the genus Scaria follows Cadena-Castafeda et al. (2019), who divided it by means of a cladistic analysis into three species groups— groups S. hamata (De Geer, 1773), S. linea- ta Bolivar 1887, S. producta Hancock, 1907—and S. laeta Giinther, 1940 without assignment to any species group. Comparative material examined.—Materials of Scaria species iden- tified by Cadena-Castafieda et al. (2019) for which photographs were available and which were used by us for comparison with the new species are included in Table 1. Museum collections acronyms used are as follows: ANSP—The Academy of Natural Sciences of Drexel University, Philadelphia, Pennsylvania, USA; BY UC—Mon- te L. Bean Life Science Museum, Brigham Young University, Provo, Utah, USA; CAUD—Coleccién de Artr6podos y otros Invertebra- dos de la Universidad Distrital Francisco José de Caldas, Bogota, Colombia; INPA—Colecao de Invertebrados do Instituto Nacional de Pesquisas da Amazonia, Manaus, Brazil; JsC—Collection Josip Skejo, Zagreb, Croatia; MNCN—Museo Natural de Ciencias Natu- rales, Madrid, Spain; MRSNT—Museo Regionale di Scienze Natu- rali di Torino, Torino, Italy; NUYW—Naturhistorisches Museum Wien, Vienna, Austria; SMDT—Staatliches Museum fiir Tierkunde, Dresden, Germany; UFAM—Colecao Zoologica Prof. Paulo Buh- rnheim, Universidade Federal do Amazonas, Manaus, Brasil. Morphological terminology.—Terminology of morphological characters follows Rehn (1904), Grant (1962), and Tumbrinck 175 (2014). Batrachideinae is a subfamily with well-defined synapo- morphies: 1) antennae usually more than 20 antennomeres, 2) anterior and mid femora with dorsal furrow, 3) female’s sper- matheca with two diverticula, 4) square-shaped paranota, 5) frontomedial projection in the form of a spine, 6) fastigium of the vertex continuously and uniformly curved and marginally projected above the compound eyes, and 7) (usually) maculated tegmina. These characters are taxonomically reliable and have already been thoroughly discussed by several authors who have also listed exceptions (Bolivar 1887, Rehn 1904, Grant 1962, Tumbrinck 2014, Tumbrinck and Skejo 2017). Division of the genus Scaria into four species groups was supported by differ- ences in the I) morphology of pronotum, II) male and female terminalia, III) head morphology, IV) general appearance of the anterior spine of pronotum, and V) body coloration. Key for identification of the species groups of genera Scaria was given in Cadena-Castanieda et al. (2019). Cadena-Castaneda et al. (2019) state that characters such as postocular stripe and tegmina macu- lation might represent homoplastic characters but are still useful in the identification key. Artificial intelligence (AI) enhancement of photographs.—An AI ser- vice (letsenhance.io) was used to enhance the photographs from iNaturalist. We used the “smart enhance” setting with 2x upscal- ing. These enhanced photographs were used to discern small de- tails, in combination with the original photographs, in order to avoid any potential visual artifacts in the upscaled photographs (Fig. 2). Al photography enhancement has proven to be a very use- ful tool in our case. We would like to note that it should be used with a measure of precaution, as the tool is not perfect, and visual artifacts in the upscaled photographs may appear. These problems are easily noted and avoided by people with experience with, in this case, animal morphology and anatomy. Results Taxonomy Family Tetrigidae Rambur, 1838 Subfamily Batrachideinae Bolivar, 1887 Tribe Batrachideini Bolivar, 1887 Genus Scaria Bolivar, 1887 Type species.—Scaria hamata (De Geer, 1773). Composition and distribution.—With this unnamed species, ge- nus Scaria currently includes 13 species. All the species inhabit South America, with only one species (S. fasciata) reaching central America (e.g., Panama and Nicaragua) (Hancock 1907, Cadena- Castaneda et al. 2019). The genus is currently divided into four species groups (see Table 1). Members of Scaria inhabit the Ama- zon rainforest and its vicinity. Generic identification of the unnamed species.—Grant (1956) listed three characters as differentiable between Scaria and newly es- tablished Rehnidium Grant, 1956: 1) morphology of female sub- genital plate, 2) smaller body size of Rehnidium members, and 3) frontal costa more protrusive in Rehnidium than in Scaria. Cade- na-Castafieda et al. (2019) added a few more characters, among them 4) Scaria is generally slenderer than Rehnidium; 5) tegmina of Scaria members are black, while they are brown in all other JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) 176 N. KASALO, M. DERANJA, K. ADZIG, R. SINDACO AND J. SKEJO Table 1. Material of Scaria species examined and organized by species groups by Cadena-Castanieda et al. (2019) containing brief in- formation on the distribution. Species Scaria (hamata) group S. boliviana Bruner, 1920 Material examined (1) NT: 192 Bolivia: La Paz: Nor Yungas, Yolosa ($16.24, W67.74) 1260 ma.s.l., 13.XII.2008. leg. S.M. Clark (BYUC); (2) 1¢ Bolivia: La Paz: Nor Yungas, Pacallo, ($16.21, W 67.79) 29.IV. 2005. leg. S.M. Distribution Bolivia (La Paz, Santa Cruz) Clark & R.L. Johnson (BYUC) S. granti Cadena- Castaneda, Mendes & Silva, 2019 S. hamata (De Geer, 1773) S. jonasi Cadena- Castaneda, Mendes & Silva, 2019 S. rafaeli Cadena- Castafieda, Mendes & (1) 14 Brasil (Cadena-Castafieda et al. 2019, fig. 24); (2) 19 Brasil (Cadena-Castafieda et al. 2019, fig. 25) Silva, 2019 Scaria (lineata) group (1) 12 Ecuador: Cachabi, leg. Rosenberg (ANSP) (photographic record (Cadena-Castaneda et al. 2019, fig. 13)); (2) 12 Colombia: Choco: Bahia Solano (N6.21, W77.40) (photographic record (Cadena- Castafieda et al. 2019, fig. 33)), (3) 1¢ (photographic record (Cadena-Castafieda et al. 2019, fig. 14)) S. fasciata Hancock, 1907 HT: 16 Brasil: Acre: Bujari, Floresta Estadual Antimary ($9.33, W68.32) 27.VII. 2016. leg. J.A. Rafael (INPA) (1) NT: 1¢ Brasil: Amazonas: Universidade Federal do Amazonas (UFAM) (S3.09, W59.97) 03.11.1979. leg. J.A. Rafael (INPA); (2) 12 Ecuador: Zamora-Chinchipe, Rio Zamora valley, El Pangui, Marali (S3.71, W78.55) 900 m a.s.]. 29.XII. 2009. leg. H. Braun, det. J. Skejo (JSC) HT: 1 Brasil: Amazonas: Tefé ($3.33, W64.69) 01.-05.XI.2016. leg. J.A. Oliveira & D.M.M. Mendes (INPA) Brazil (Acre state: Bujari, Floresta Estadual Antimary) Brazil (Amazonas state); Ecuador Brazil (Amazonas state, Tefé) Brazil (Amazonas state and Rond6énia state) Colombia; Ecuador; Panama; Nicaragua (1) 12 Brasil: Rond6énia: Candeias do Jamari, Usina Hidrelétrica de Samuel ($8.95, W63.18) S. ferruginea Hancock, 1909 17.VHI.2016. leg. D.M.M. Mendes, EF. Xavier F°, A.A. Agudelo, & J.A. Rafael (INPA); (2) 19 Bolivia: La Paz: Parque Nacional Madidi, (photographic record (Cadena-Castaneda et al. 2019)); (3) 19 Brazil (Rond6énia state); Colombia Colombia: Vaupés: Mitu (N1.27, W70.22) (photographic record (Cadena-Castaneda et al. 2019)) S. lineata Bolivar, 1887 (1) LT: 1¢ Peru: Alto Amazonas ($5.56, W76.00) (MNCN); (2) PLT: 19 Peru: Alto Amazonas ($5.56, W76.00) (MNCN) (photographic record (Cadena-Castaneda et al. 2019, figs 15, 16)) Ecuador (Pastaza Province); Peru (Department of Loreto) (1) HT: 19 Peru: Junin, Puerto Bermudez, Rio Pichis, 12-19.VI.1920 (ANSP) (photographic record: Cadena-Castaneda et al. 2019, fig. 17) (wrongly cited ‘male’ in OSF); (2) 19 Bolivia: PN Madidi (photographic record (Cadena-Castafieda et al. 2019, fig. 34)); (3) 12 Peru, PN. Manu, Pantiacolla rainforest (photographic record: http://orthoptera.speciesfile.org/Common/basic/ShowImage.aspx?Tax S. veruta (Grant, 1956) Peru (Rio Pichis, Puerto Bermudez) onNameID=1100469&lmagelD=203222) Scaria (producta) group S. maculata Giglio-Tos, 1898 LT: 14 Ecuador: Valle de Santiago ($3.53, W78.46) (MRSNT) (1) 12: Peru: Loreto, Picuroyacu, 2013, (photographic record (Cadena-Castafieda et al. 2019, fig. 32)); (2) 12: Colombia: Putumayo: Mocoa (N1.15, W76.64) 500 m a.s.l. 2017. (Photographic record S. producta Hancock, 1907 Ecuador (Valle de Santiago) Colombia; Ecuador; Peru (Cadena-Castaneda et al. 2019, fig. 32)) Scaria (laeta) group (1) LT: 14 Brasil: Amazonas: Sao Paulo de Olivenga, southern banks of upper Amazonas, leg. S. & I. Waehner (SMTD); (2) PLT: 19 Brazil, Amazonas: Sao Paulo de Olivenca, mouth of Rio Javary S. laeta (S3.76, W69.09) leg. S. & I. Waehner (SMTD); (3) PLT: 14 Brasil: Amazonas: Sao Paulo de Olivenga, Colombia; Ecuador; southern banks of upper Amazonas, leg. S. & I. Waehner (SMTD); (4) PLT: 19 Brasil: Amazonas: Sao Paulo de Olivenga (S3.38, W69.07) 65 mazs.l., leg. S. Waehner (NMW); (5) 1° Brasil: Amazonas: Tefé (photographic record (Cadena-Castafieda et al. 2019, fig. 31)); (6) 14 (photographic record (Cadena- Castaneda et al. 2019, fig. 8)); (7) 12 (photographic record (Cadena-Castanieda et al. 2019, fig. 9)) Giinther, 1940 genera of Batrachideinae; and 6) hind femora sulcated in Scaria, unlike carinated in Rehnidium. We identify this species as Scaria Bolivar, 1887 because it shows the following characters typical of the genus: 1) median carina of the pronotum projected above the head anteriorly in the form of a well-developed spine (similar to Rehnidium Grant, 1956 in which the spine is less pronounced); 2) slender (slim) and elongated body (differing it from robust mem- bers of the genus Rehnidium Grant, 1956); 3) black basal color of Peru tegmina (not brown as in Rehnidium); and 4) flat pronotum (not roof-like as in Rehnidium). The two individuals of the newly reported species are macrop- terous and macropronotal, pronotum being longer than hind femora. Characters that are present in all Scaria species known so far, but at first seem to be absent in the unnamed one, are 1) ab- sence of clearly visible pale colored spot at posterior part of teg- men and 2) a lack of dark lateral stripe on the pronotum. The JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) N. KASALO, M. DERANJA, K. ADZIC, R. SINDACO AND J. SKEJO tegminal spot might be present in the reported specimens, but it is simply covered by a wide yellow-colored stripe that lies in the middle part of tegmen. This is the first Scaria species with medial stripe on tegmina (for comparison, S. laeta Giinther, 1940 has the yellow stripe in the ventral portion of the tegmen, while S. jonasi Cadena-Castaneda et al. 2019 has it on the dorsal portion of teg- mina). Arguments for not identifying the species as Rehnidium are 1) members of Rehnidium always have a brown base color of teg- mina and 2) members of the genus Rehnidium have short and wide ovipositor valves (Cadena-Castaneda and Cardona 2015, Cadena- Castaneda et al. 2019). Scaria sp. Material examined.—Perru ¢1 M, 1 F; Department of Amazonas: Bon- gara Province: Peroles near Yambrasbamba, mountain rainforest belonging to the Peruvian Yungas biogeographic ecoregion; 5.67°S, 77.92°W; 1905 ma.s.l.; 19 August 2008; R. Sindaco leg.; photographs only, available on iNaturalist (https://www.inaturalist.org/observa- tions/9968031), supplemented by Figures in this publication. Habitat.—The specimens were observed and photographed on the ground inside a well-preserved patch of forest with muddy base covered by abundant leaf litter; tree trunks were covered by mosses, rich epiphytic vegetation (many Bromeliaceae), and arboreal ferns. Specific traits. —The main differences between Scaria sp. and the five other morphologically similar species that occur in the region are listed in Table 2. The unnamed species is easily distinguished from other Scaria species by the following set of characters: 1) vertex wider than in any other known Scaria species, 2) small apical teeth on mid femora (in almost all the other species, teeth are larger), 3) longer fore and mid femora (length/width ratio of 6 or more) than in any other species, 4) generally stouter appearance than any other Scaria species, 5) yellow stripe placed medially on tegmina, and 6) distinctive coloration pattern of pronotum. Concerning the key to Scaria species by Cadena-Castaneda et al. (2019), our unnamed species shows a unique combination of characters not present in any other known species, namely yellowish face, eyes 177 projected above the dorsalmost level of tegmina, absence of dorsal midline (coloration), lightly colored stripe covering the median part of tegmina, and absence of stripe on the upper half of the lateral margin of pronotum. The only Scaria species similar to this unnamed one is Scaria veruta (Grant, 1956), which also has longer fore and mid femora. Morphological description.—General characters and coloration: Relatively slender body, slightly stouter than other Scaria species described so far. Body smooth, without warts or dorsal projec- tions. Coloration pattern of the entire body homogeneous, with interchanging black and yellow coloration. Yellow coloration var- ying from dark to paler yellow (close to white coloration) in some parts of the body. Antennae black, sometimes with a paler tip. Head exhibiting similar coloration and texture to that of the rest of the body, with black stripe behind the eyes. Compound eyes pale at the top, otherwise black. Carinae of pronotum (interhumeral carinae, external lateral carinae, internal lateral carinae, prozonal carinae, and median carina) mostly yellow to light yellow. Fore and mid femora black with all carinae yellow; fore and mid tibiae following the same pattern. Hind femora bearing a yellow to light yellowish stripe in the mid part. Front and mid tarsi dark; hind tarsi lighter but with dark coloration on the first segment ventrally. Tegmen black with a yellow longitudinal stripe in the middle cov- ering more than three quarters of the area of the tegmen. Head: (Fig. 2A, E) Antennae with 20 antennomeres. Scapus oval in cross-section. Short axis of the scapus 1.5 times wider than second antennomere (pedicel); long axis 2 times wider than sec- ond antennomere. Second antennomere 1.5 times wider than the third and all other antennomeres. Basal antennomeres from 3” to 7, central antennomeres from 8" to 14", preapical antenno- meres from 15" to 17", and apical antennomeres final three seg- ments (18" to 20") reduced. Pale rings in joints visible in central segments. In fronto-lateral view: Frontal costa bifurcates above the middle of the compound eyes (Fig. 5). Head below level of pronotum (head in contact with the underside of the anterior side of pronotum). Lateral ocelli barely above mid-level of a com- pound eye. Fastigium verticis barely below level of dorsal margin of a compound eye. Frontal costa bulging for half the length of Table 2. Tabular comparison of Scaria sp. to five other species that are morphologically similar or found in the same area. Scaria sp. S. hamata S. lineata S. maculata _ S. ferruginea S. veruta fore femur length/width (height) ratio 6 3.9-4.5 4.7-5.1 5xh 4.65-4.85 6.25 relation between lateral and humero-apical carinae parallel almost touch almost touch parallel almost touch almost touch PRs ; : absent or very convexity in fronto-lateral area of pronotum noticeable slight absent absent k absent wea a rounded, pointed, rounded, pointed, rounded, bulbous, tarsal pulvilli . ; j bulbous triangular bulbous triangular triangular rounded apical teeth of mid femora small large medium-large small-medium medium-large medium ‘ ; 1 (male), 1.7 width of vertex to compound eye length (dorsal view) LS kS 1 (male) 1.4 1.2 (female) (female) ; sometimes . PM elevation present present absent present slight present humeral angles rounded rounded rounded _ slightly angled slightly angled rounded rising of pronotum after PM towards anterior spine very slight slight pronounced pronounced pronounced pronounced ovipositor valve length long medium medium-long N/A short medium-long barel pronotum length in comparison with hind knee surpassing surpassing surpassing surpassing surpassing e surpassing : absent or very post ocular stripe present present present absent k present wea tegminal coloration wide band spot no coloration no coloration spot spot JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) 178 ia = N. KASALO, M. DERANJA, K. ADZIG, R. SINDACO AND J. SKEJO Fig. 2. Example of AI enhancement of Scaria sp. body details compared to original photography in the background. A. Upper part of the head, showing an eye, a scapus, and a pedicel; B. Ovipositor; C. Hind tarsus; D. Ventral and tegminal sinuses; E. Bottom part of the head, showing the mouthparts. Photo credit: Roberto Sindaco. scapus. Pedipalps white. Transverse carinae concave. Fossulae deep and pronounced. Median carina absent. In dorsal view: Vertex the same width as compound eye (male) or 1.7 times as wide as com- pound eye (female). Eyes of a bulbous kidney shape. Frontal costa not straight after bifurcation (see Fig. 4A). Pronotum: Frontal view is not seen in the photographs. In lateral view: Pointy short frontomedial (anterior spine) present in the anterior margin. Prozonal carina visible, yellow in coloration. Sulci visible, dark in coloration. Humeroapical carina connected with external lateral carina, both yellow in coloration. Posterior margin of the lateral lobe yellow/pale with ventral sinus more ob- tuse than tegminal sinus. Pronotal disc flat; median carina flat ex- cept on the places of promedial and first metamedial projections. Pronotum reaching far beyond hind knees. Extralateral carina not visible, but a yellow spot in its place. Infrascapular area virtu- ally non-existent. In dorsal view: Pronotum covering the whole abdomen. Pronotal apex surpassing hind femora. Coloration of pronotum similar to rest of body. promedial projection with char- acteristic yellow coloration. Prozonal carinae parallel. Humeral angles slender, oblique. Pronotal process bearing yellow to pale yellowish x-shaped mark. Median carina present, clearly visible thanks to the contrast in coloration, but rather flat. Internal lateral carina yellow to pale yellow and clearly visible. Posterior margin of pronotum truncated. Interhumeral carina absent. Characteris- tic yellow line visible in the area where an interhumeral carina is usually present. Wings: Macropterous specimen. Wings (alae) well developed, visibly longer than pronotum, black with white anterior edge. Teg- mina present; reaching coxa of hind legs; black with thick yellow medial stripe from anterior to posterior part of tegmen. Legs: Fore legs: Femora and tibiae smooth, without teeth. Yellow and black stripes following visually unperceivable cari- nae on femora. Fore femora 6 times longer than wide. Yellow and black stripes present on tibiae as well. Tarsi two segmented. Proximal segment much shorter than distal. Mid legs: Femora and tibiae smooth except for the apical (genicular) teeth pre- sent and clearly visible in distal part of femora. Yellow and black stripes, which are 6 times longer than wide, following virtually non-existent carinae on femora. Yellow and black stripes present on tibiae. Tarsi two segmented, proximal segment much shorter than distal. Hind legs: Femora 3.6 times longer than wide. Dor- sal margin with minuscule teeth along dorsal margin. Genicular and antigenicular teeth clearly visible, but small. Ventral margin smooth. Inner external area of hind femora with a few transverse ridges of yellow color. In the mid length of femur, a transverse yellow to pale yellow band is present. Tibiae yellow in dorsal, black in ventral part, with recognizable miniscule teeth on dorsal margins with few larger, but still tiny, teeth. Third segment of JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) N. KASALO, M. DERANJA, K. ADZIC, R. SINDACO AND J. SKEJO a tes . see , y 7. By ag a Fig. 3. The picture of the habitat taken by R. Sindaco during the trip to Peru during which the pictures of Scaria sp. were taken (Homo sapiens (Alberto Venchi) for scale). arsus (Fig. 2C) 1.4 times longer than proximal segment. Tarsal pulvilli orbicular, first and second of same length, third ( = dis- tal) 1.4 times longer than first two. Proximal segment of tarsus in dorsal part yellow to light yellowish; in ventral part black, but tarsal pulvillus pale. Sexual dimorphism.—This unnamed species exhibits marked sex- ual dimorphism in general appearance of sexes, vertex width, pronotum stature, and coloration. Some of the observed dif- ferences could be due to the limited sample, i.e., we have ex- amined photographs of only one male and one female, but we nonetheless decide to discuss them as they could prove to be useful in the future. Vertex: Ratio of vertex width and width of compound eye in dorsal view much higher in female (1.7) than in male (1). Pronotum: Ratio of width between prozonal carinas and the width between humoral angles are equal in both sexes. However, the ratio of the length of the pronotum and the widest width between the humeral angles is much less in the female (5.15) than in the male (5.7). Coloration: Observed pattern of coloration is almost identical in both sexes, with varying degrees of color saturation in certain areas. Legs in male appear more saturated than in female. Pronotum of female appears more saturated than that of male. Fig. 4. Living female of Scaria sp. in A. Dorsal view; B. Lateral view. Photo credit: Roberto Sindaco. Limits of the description from photographs.—Since some angles are missing from the pictures, certain characters were described incompletely or were not described at all. We must stress, once again, that those characters do not limit us in concluding that this is a new species, but could limit the comparison with future speci- mens or photographs. The following is a list of characters seen only in frontal and ventral views, which should, for detailed de- scription, be examined in a laboratory environment when a physi- cal specimen is collected. Frontal view: level of bifurcation of the frontal costa; scutellum width; position of the antennal grooves; distance between the antennal grooves; position of lateral ocellus; shape of the vertex. Ventral view: shape of thoracal and abdominal sternites (incl. sternomentum and subgenital plate). Measurements.—No specific measurements can be given due to lack of physical specimens, but some specific proportions can be calculated from photographs (see Table 2). Discussion On the taxonomy of Scaria Specimens of Scaria sp. that we report here are somewhat simi- lar to the holotype of Scaria veruta (originally placed in Rehnidium JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) Fig. 5. Living female of Scaria sp. in frontolateral view. Photo cred- it: Roberto Sindaco. Grant, 1956) (Grant 1956, Cadena-Castanieda et al. 2019, Silva et al. 2021). The lack of a lateral stripe on the pronotum of Scaria sp. might be caused by the fact that our species has a multicolored pronotum on which the stripe might simply be difficult to notice and distinguish. A similar case has been observed in a living speci- men of S. hamata (De Geer, 1773) reported by Cadena-Castaneda et al. (2019: fig. 36). In the key given by Cadena-Castaneda et al. (2019), Scaria veruta can be distinguished from other species by tegmen with an ovoid subapical spot. However, the key then for- wards a reader to the previously mentioned figure 17, a specimen of S. veruta with yellowish ventral line on the tegmina. Also, figure 34 shows a living specimen of S. veruta with tegminal sinus and tarsal pulvilli different from the ones on specimen from figure 17. This brings us to question whether all these specimens belong to the same species. Describing species from photographs On the nature of the name The nature of the name is an important point deserving to be separately discussed. Despite the existing rules of nomenclature provided by the ICZN, we did not manage to “legally” name this new species. The name is one of the first steps in investigating any species, learning about its behavior, habitat, and distribu- tion. Article 72.5.6. of the ICZN states, “In the case of a nominal species-group taxon based on an illustration or description, or on a bib- liographic reference to an illustration or description, the name-bearing type is the specimen or specimens illustrated or described (and not the N. KASALO, M. DERANJA, K. ADZIG, R. SINDACO AND J. SKEJO Bae