Research Article |
Corresponding author: Niko Kasalo ( niko.kasalo5@gmail.com ) Corresponding author: Josip Skejo ( skejo.josip@gmail.com ) Academic editor: Maria-Marta Cigliano
© 2021 Niko Kasalo, Maks Deranja, Karmela Adžić, Roberto Sindaco, Josip Skejo.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Kasalo N, Deranja M, Adžić 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
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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 nomenclatural actions, a new species of the Batrachidein pygmy grasshoppers belonging to the genus Scaria Bolívar, 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.
Amazon, conservation, new species, Orthoptera, Peruvian Yungas, photography-based taxonomy, pygmy grasshopper, the ICZN
Describing species from photographs or illustrations is not a common practice, but it has been done a number of times (
While
In this paper, we describe an unnamed species of a pygmy grasshopper (family Tetrigidae, subfamily Batrachideinae Bolívar, 1887), belonging to the tribe Batrachideini and genus Scaria Bolívar, 1887 (according to
Locality information.—The unnamed Scaria species is, for now, known only from a single locality in Peru. Peroles, near Yambrasbamba 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,
Taxonomy.—Taxonomy follows Orthoptera Species File (OSF;
Comparative material examined.—Materials of Scaria species identified by
Material of Scaria species examined and organized by species groups by
Species | Material examined | Distribution |
---|---|---|
Scaria (hamata) group | ||
S. boliviana Bruner, 1920 | (1) NT: 1♀ Bolivia: La Paz: Nor Yungas, Yolosa (S16.24, W67.74) 1260 m a.s.l., 13.XII.2008. leg. S.M. Clark (BYUC); (2) 1♂ Bolivia: La Paz: Nor Yungas, Pacallo, (S16.21, W 67.79) 29.IV. 2005. leg. S.M. Clark & R.L. Johnson (BYUC) | Bolivia (La Paz, Santa Cruz) |
S. granti Cadena-Castañeda, Mendes & Silva, 2019 | HT: 1♂ Brasil: Acre: Bujari, Floresta Estadual Antimary (S9.33, W68.32) 27.VII. 2016. leg. J.A. Rafael (INPA) | Brazil (Acre state: Bujari, Floresta Estadual Antimary) |
S. hamata (De Geer, 1773) | (1) NT: 1♂ Brasil: Amazonas: Universidade Federal do Amazonas (UFAM) (S3.09, W59.97) 03.II.1979. leg. J.A. Rafael (INPA); (2) 1♀ Ecuador: Zamora-Chinchipe, Río Zamora valley, El Pangui, Maralí (S3.71, W78.55) 900 m a.s.l. 29.XII. 2009. leg. H. Braun, det. J. Skejo (JSC) | Brazil (Amazonas state); Ecuador |
S. jonasi Cadena-Castañeda, Mendes & Silva, 2019 | HT: 1♂ Brasil: Amazonas: Tefé (S3.33, W64.69) 01.–05.XI.2016. leg. J.A. Oliveira & D.M.M. Mendes (INPA) | Brazil (Amazonas state, Tefé) |
S. rafaeli Cadena-Castañeda, Mendes & Silva, 2019 | (1) 1♂ Brasil ( |
Brazil (Amazonas state and Rondônia state) |
Scaria (lineata) group | ||
S. fasciata Hancock, 1907 | (1) 1♀ Ecuador: Cachabi, leg. Rosenberg (ANSP) (photographic record ( |
Colombia; Ecuador; Panama; Nicaragua |
S. ferruginea Hancock, 1909 | (1) 1♀ Brasil: Rondônia: Candeias do Jamari, Usina Hidrelétrica de Samuel (S8.95, W63.18) 17.VIII.2016. leg. D.M.M. Mendes, F.F. Xavier F°, A.A. Agudelo, & J.A. Rafael (INPA); (2) 1♀ Bolivia: La Paz: Parque Nacional Madidi, (photographic record ( |
Brazil (Rondônia state); Colombia |
S. lineata Bolívar, 1887 | (1) LT: 1♂ Peru: Alto Amazonas (S5.56, W76.00) (MNCN); (2) PLT: 1♀ Peru: Alto Amazonas (S5.56, W76.00) (MNCN) (photographic record ( |
Ecuador (Pastaza Province); Peru (Department of Loreto) |
S. veruta (Grant, 1956) |
(1) HT: 1♀ Peru: Junín, Puerto Bermudez, Río Pichis, 12–19.VII.1920 (ANSP) (photographic record: |
Peru (Río Pichis, Puerto Bermúdez) |
Scaria (producta) group | ||
S. maculata Giglio-Tos, 1898 | LT: 1♂ Ecuador: Valle de Santiago (S3.53, W78.46) (MRSNT) | Ecuador (Valle de Santiago) |
S. producta Hancock, 1907 | (1) 1♂: Peru: Loreto, Picuroyacu, 2013, (photographic record ( |
Colombia; Ecuador; Peru |
Scaria (laeta) group | ||
S. laeta Günther, 1940 | (1) LT: 1♂ Brasil: Amazonas: São Paulo de Olivença, southern banks of upper Amazonas, leg. S. & I. Waehner (SMTD); (2) PLT: 1♀ Brazil, Amazonas: São Paulo de Olivença, mouth of Rio Javary (S3.76, W69.09) leg. S. & I. Waehner (SMTD); (3) PLT: 1♂ Brasil: Amazonas: São Paulo de Olivença, southern banks of upper Amazonas, leg. S. & I. Waehner (SMTD); (4) PLT: 1♀ Brasil: Amazonas: São Paulo de Olivença (S3.38, W69.07) 65 m a.s.l., leg. S. Waehner (NMW); (5) 1♀ Brasil: Amazonas: Tefé (photographic record ( |
Colombia; Ecuador; Peru |
Morphological terminology.—Terminology of morphological characters follows
Artificial intelligence (AI) enhancement of photographs.—An AI service (letsenhance.io) was used to enhance the photographs from iNaturalist. We used the “smart enhance” setting with 2× upscaling. These enhanced photographs were used to discern small details, in combination with the original photographs, in order to avoid any potential visual artifacts in the upscaled photographs (Fig.
Family Tetrigidae Rambur, 1838
Subfamily Batrachideinae Bolívar, 1887
Tribe Batrachideini Bolívar, 1887
Scaria hamata (De Geer, 1773).
With this unnamed species, genus 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) (
The two individuals of the newly reported species are macropterous 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) absence of clearly visible pale colored spot at posterior part of tegmen and 2) a lack of dark lateral stripe on the pronotum. The 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 Günther, 1940 has the yellow stripe in the ventral portion of the tegmen, while S. jonasi
Peru •1 M, 1 F; Department of Amazonas: Bongará Province: Peroles near Yambrasbamba, mountain rainforest belonging to the Peruvian Yungas biogeographic ecoregion; 5.67°S, 77.92°W; 1905 m a.s.l.; 19 August 2008; R. Sindaco leg.; photographs only, available on iNaturalist (https://www.inaturalist.org/observations/9968031), supplemented by Figures in this publication.
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.
The main differences between Scaria sp. and the five other morphologically similar species that occur in the region are listed in Table
General characters and coloration: Relatively slender body, slightly stouter than other Scaria species described so far. Body smooth, without warts or dorsal projections. Coloration pattern of the entire body homogeneous, with interchanging black and yellow coloration. Yellow coloration varying 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 covering more than three quarters of the area of the tegmen.
Head: (Fig.
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 obtuse than tegminal sinus. Pronotal disc flat; median carina flat except 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 virtually 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 characteristic 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. Characteristic 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. Tegmina 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 carinae 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 present 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. Dorsal 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 arsus (Fig.
This unnamed species exhibits marked sexual dimorphism in general appearance of sexes, vertex width, pronotum stature, and coloration. Some of the observed differences could be due to the limited sample, i.e., we have examined 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.
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 specimens or photographs. The following is a list of characters seen only in frontal and ventral views, which should, for detailed description, be examined in a laboratory environment when a physical 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).
No specific measurements can be given due to lack of physical specimens, but some specific proportions can be calculated from photographs (see Table
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 | 5.1 | 4.65–4.85 | 6.25 |
relation between lateral and humero-apical carinae | parallel | almost touch | almost touch | parallel | almost touch | almost touch |
convexity in fronto-lateral area of pronotum | noticeable | slight | absent | absent | absent or very weak | absent |
tarsal pulvilli | rounded, bulbous | pointed, triangular | rounded, bulbous | pointed, triangular | rounded, triangular | bulbous, rounded |
apical teeth of mid femora | small | large | medium-large | small-medium | medium-large | medium |
width of vertex to compound eye length (dorsal view) | 1 (male), 1.7 (female) | 1.3 | 1.3 | 1 (male) | 1.4 | 1.2 (female) |
PM elevation | present | present | absent | present | sometimes present | slight |
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 |
pronotum length in comparison with hind knee | surpassing | surpassing | surpassing | surpassing | surpassing | barely surpassing |
post ocular stripe | present | present | present | absent | absent or very weak | present |
tegminal coloration | wide band | spot | no coloration | no coloration | spot | spot |
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.
Specimens of Scaria sp. that we report here are somewhat similar to the holotype of Scaria veruta (originally placed in Rehnidium Grant, 1956) (
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 distribution. 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 bibliographic reference to an illustration or description, the name-bearing type is the specimen or specimens illustrated or described (and not the illustration or description itself).” Article 73.1.4. states, “Designation of an illustration of a single specimen as a holotype is to be treated as designation of the specimen illustrated; the fact that the specimen no longer exists or cannot be traced does not of itself invalidate the designation.” Recommendation 73B of the ICZN states, “An author should designate as holotype a specimen actually studied by him or her, not a specimen known to the author only from descriptions or illustrations in the literature” (ICZN 1999). It is clear that, despite the recommendation that new species should be based on physical specimens, it is not forbidden to describe a species from a photograph, i.e., from the specimen a photograph represents, if there is reason to do so. As 73B is ‘only’ a recommendation, it has provided room for well-documented exceptions from physically collected species (
The ICZN anticipated descriptions of typeless species. Recommendation 73G asks the authors to provide an explanation for why they lack physical types, and this is asked in order to avoid “immediate science,” i.e., fast typeless descriptions of taxa found in online photographs but for which physical types could easily be provided. The ICZN asks as follows: “An author should provide detailed reasoning why at least one preserved specimen, whether a complete individual organism or a part of such an individual, was not used as the name-bearing type for the new taxon and why the formal naming of the taxon is needed at a point in time when no preserved name-bearing type will be available” (ICZN 1999).
A name of any kind would make referring to this species easier in subsequent publications. A proper binomial name would carry taxonomic information. A nomenclatural valid name would allow species systematization in databases of all kinds.
The ICZN states that “a holotype is the single specimen upon which a new nominal species-group taxon is based in the original publication” (ICZN 1999). The holotype is only one because numerous problems existed with the syntypes, such as more than one species being contained within a single type series (
Another problem is the inaccessibility of certain holotypes, either in museums (
The taxonomic community seems content with the idea that the holotype merely exists somewhere but does not clearly mandate the ways of their preservation, accessibility, or eventual replacement due to inevitable degradation. This is excusable if the function of the holotype is to provide the basis for a name (which it is) but is equivalent to institutionalized malpractice if the holotype is supposed to be accessible at any moment (which it is not). If the designated holotype or the depiction of it represents sufficient evidence to differentiate the named taxon from the others, its function is fulfilled. The missing data can be acquired in subsequent research and published separately. Publishing a species name does not entail providing complete and non-editable information about the species.
The decision to try to describe a new species from photographs (iNaturalist) was made after we found no similar pygmy grasshopper in the literature. This nameless Scaria was recorded only once in 2008 in a severely deforested area. The fact that we made no attempt to physically find it, either by ourselves or with help from potential colleagues in Peru, is one of the criticisms we received, and it certainly warrants an explanation. After examining the photographs of the specimens and concluding that they represent a currently unknown species, we felt obliged to add this species to the globally accessible fund of knowledge, which entails naming it according to the accepted rules. As this was all we set out to do and the data available to us allowed for delimiting the new species from its congeners, there is no need for additional research at this stage.
Plenty of papers that present only basic information, i.e., without DNA sequences (
Field work is a necessary part of research, and we will, in time, take steps to either visit the locality or have it visited by somebody else. Considering the cost of this endeavor, the time it takes to forge a robust relationship with local researchers, and the restrictions imposed by the pandemic, this will happen later rather than sooner.
Incidentally, research concerning the genus Scaria in an area that encompasses the area in which the nameless Scaria lives was recently conducted by
The existence of protected areas where collecting specimens is prohibited and even touristic visits are heavily regulated is very positive, and governments around the world should be urged to designate more such areas for obvious reasons. However, those strict limitations can impede research to the point that many expeditions of a smaller scope will simply not be undertaken. It is impossible to acquire permits for those areas, as the process is arduous and (understandably) discouraging (
Even though the name carries a certain significance by itself, it also has one strictly practical value. Laws are written in such a way as to direct conservation efforts towards entities bearing a certain name, and not towards the species itself. Changing the name can impact the conservation status of a species regardless of its actual status (IUCN SSC 2012, ICZN 2021).
From the available data, it is impossible to make less than speculative statements about the status of the nameless Scaria, and we will refrain from doing so. It does not have to be extinct or even endangered for these arguments to have weight. Every species should be named if the naming can be done legitimately. This allows the law to recognize it and take appropriate action. The name can be changed in cases of fresh discoveries, so there is no apparent reason to circumvent rules in order to disallow a name and, in doing so, make further research on the species more difficult while robbing the area’s checklist of a new species. What is discussed here is principle, not the significance of a single species.
There is a nameless new Scaria species from Peru, clearly distinguishable from other species of the genus. All of the species of that genus are clearly differentiable by their morphology and coloration, which are visible in photographs. The ICZN allows naming species from photographs if there is reason to do so; however, we did not manage to name this nameless species. We do not propose that photograph-based descriptions become the norm, only that in certain cases and for certain taxa this can be beneficial. We understand how important the physical holotype is, but because of numerous issues, we did not manage to collect one.
By the time a scientist collects, examines, and describes one new species, several more have gone extinct (
We would like to acknowledge iNaturalist for providing an excellent platform for encouraging citizen science. This service allows people without formal training to participate in collecting valuable data that can help researchers around the globe. This paper is only a single example of the possibilities that are created by services such as iNaturalist. Thanks to Daniela Santos Silva for clarifying whether figure 17. in