A new species of Zetekella Drake from Ecuador with comments on Zetekella and Minitingis Barber (Heteroptera, Tingidae)

Abstract Zetekella and Minitingis (Heteroptera, Tingidae) are morphologically similar genera, each comprising two species. The latter was already considered a junior synonym of the former, but was revalidated on the basis of the number of cephalic spines, projections on the paranotal edge, length of the rostrum, presence of an abdominal groove and distributional pattern. Here, a new species of Zetekella is described from Ecuador, the diagnoses for both genera reassessed, new records for Z.pulla and Z.zeteki reported, and a key to the species of both genera provided.


Introduction
Zetekella Drake is composed of two species, Z. zeteki Drake, 1944 andZ. pulla Drake &Plaumann, 1956. After Z. pulla was described, the generic diagnosis was redefined, as follows: head moderately long to long, armed with five spines, bucculae open in front and slightly projected forward, and "rostrum extremely long, extending on venter" (Drake and Plaumann 1956). No macropterous forms are known for this genus, but other characters, such as the proportions of the antennal segments, often have been used in taxonomic studies of the Tingidae (excluding Vianadinae).
Zetekella was considered the senior synonym of Minitingis Barber by Drake and Ruhoff (1960) without further consideration of morphological characters or generic diagnoses. This genus was originally proposed to hold Minitingis minusculus Barber, 1954 on the basis of the number of pronotal carinae and the lateral acute processes of the paranota. However, the genus was compared with Phatnoma, rather than Zetekella, and the remarkable paranotal acute processes were found to vary by the same author (Barber 1954). Froeschner (1968) reinstated Minitingis, described a new species of the genus, and reaffirmed the generic status based on morphological characters and distributional patterns. According to Froeschner (1968), Minitingis could be distinguished by the presence of seven cephalic spines, the occipital pair being short and obliquely elevated, and the rostrum reaching the second abdominal segment. The paranotal development and the abdominal groove were also mentioned as diagnostic features of the genus (Froeschner 1968). Both M. minusculus and M. elsae Froeschner, 1968 are from the West Indies, whereas the known species of Zetekella are from Panama and Brazil. This distribution represents different zoogeographical zones and, therefore, corroborates the hypothesis of two genera (Froeschner 1968).
In this paper, we describe a new species of Zetekella from Ecuador, report two new records for Z. pulla and a new country record for Z. zeteki, and re-evaluate the diagnostic characters of both genera.

Material studied
The specimen here described was collected in a Berlese trap and had its abdomen removed for DNA extraction. The fixation method of the specimen is unknown, and it was preserved in 75% alcohol before the abdomen was removed and the specimen mounted. The specimen was point-mounted on the left side instead of the right side, to preserve two of its legs that accidentally had come in contact with the glue during the mounting process.
Holotypes of all species (except M. minusculus) were studied. For M. minusculus, a six-specimen series of paratypes was analyzed. All type material was examined at the National Museum of Natural History (USNM), in Washington, D.C., USA. Fifteen specimens of Z. pulla from the Museu de Zoologia da Universidade de São Paulo, Brazil, were also studied. The remaining specimens are housed in the first author's personal collection.

Species descriptions
Measurements of the holotype were taken from photos using ImageJ and are given in millimeters. Terminology follows the specialized literature Davis 1960, Drake andRuhoff 1965). The taxonomic act here treated was registered in Zoobank (Pyle and Michel 2008).

Images
Photos were taken with a camera attached to a stereoscope and treated in GIMP. Plates were composed in Inkscape. The holotype photos of Z. pulla, Z. zeteki, and M. minusculus were kindly provided by Thomas Henry. Dorsal habitus and labels of the holotypes, voucher specimens for the new records, the two paratypes, and lateral and ventral views of the holotype of the new species were photographed and made available at Figshare.

Keys
The keys to Minitingis and Zetekella species provided by Froeschner (1996) were merged, adapted and updated to include new species and new findings.

Occurrence data
Geographic coordinates, when not available on the specimen labels, were obtained using Google Earth. The map was built using SimpleMappr (Shorthouse 2010). This map includes a layer with the Biodiversity Hotspots (sensu Conservation International; Mittermeier et al. 2004). Additionally, a spreadsheet containing occurrence data extracted from specimen labels was made available at Zenodo; the spreadsheet is organized by specimens and their unique identifiers, when available.  Diagnosis. Body dark brown to blackish; cephalic spines long and thin; anterior edge of paranota not reaching the eyes; discoidal area biseriate and subcostal area irregularly quadriseriate.

Zetekella henryi
Description. Body oval; mostly dark brown, or blackish; collar, paranota, and lateral edge of costal area and hemelytral membrane white; tip of cephalic spines, scape and pedicel light brown (basi-and distiflagellomere missing); occipital spines lighter in color.
Head with numerous, small, curved hairs and seven spines: clypeal pair non-erect; jugal spine slightly erect; frontal pair divergent; occipital pair short, strongly divergent; frontal and occipital pairs erect. Antenniferous processes spine-like, projected forward, subequal to scape in size. Scape slightly longer than pedicel, basi-and distiflagellomere missing. Interocular distance almost three times width of eye. Rostrum light brown, surpassing posterior margin of metanotum. Bucculae white, areolate; open in front, with an acutely projected antero-inferior edge; widely open posteriorly, width same as anterior region.
Pronotum mostly flat, posterior projection absent, leaving small portion of scutellum exposed. Median carinae whitish, uniseriate, composed of small cells, extending throughout pronotum. Collar biseriate and slightly elevated. Paranota slightly reflexed, broad, with four cells at widest part; anterior edge not reaching eyes. Sternal membranes whitish, areolate, uniseriate, and concave. Hemelytra ovate, inner border conspicuously concave posteriorly; clavus large, 2-seriate at widest part, inner vein straight, outer edge convex; discoidal area biseriate; cubitus whitish posteriorly after R+M junction; radius-media (R+M) white for most of length, raised, stout; subcostal area mostly 3-seriate, four rows of areolae at widest part; costal area wide, with as many as six rows of areolae, widening posteriorly; membrane shortened (specimen brachypterous); hypocosta dark brown, areolate anteriorly, but light brown, rim-like for most of length, ending at membrane. Scent-gland opening round, auricular-like, dark. Legs light brown, coxae and trochanters stout; longer, spine-like setae at posterior edge of tibiae; second tarsi long and slender. Claws long, slender, well developed.
Pygophore conspicuously narrower than abdomen; dorsal rim strongly curved, almost sinuous, forming small depressions laterally and dorsally. Paramere stout at base, abruptly but consistently narrowing to very slender tip, pronounced elbow at base.
Remarks. Of the three known species of Zetekella, Z. henryi sp. n. is more morphologically similar to Z. zeteki because of the broader paranota and hemelytra, and the long clypeal, jugal and frontal cephalic spines. It differs from Z. zeteki by the thinner cephalic spines, the anterior edge of paranota not reaching the eyes, the narrower discoidal and subcostal area, and by its color pattern.
Etymology. This species is named after the outstanding heteropterist and dear friend Thomas Henry, on the occasion of his 70 th birthday and his remarkable career and countless contributions to the study of Heteroptera.  Spreadsheet containing label information of all studied specimens and their respective unique identifiers

Discussion
Zetekella henryi sp. n. is described based on morphological differences in characters that have been commonly used to delimit species within Tingidae. The new species resembles Z. zeteki, but differs from it by the color pattern, paranota, and discoidal and subcostal areas of the hemelytra. Additionally, the shorter rostrum and shape of the scent gland allies these two species with Z. pulla. In addition to the description of a new species of Zetekella, a macropterous specimen of Z. pulla was found and is illustrated. All characters, except the hemelytral membrane, remain virtually the same between the macropterous and brachypterous specimens. Only brachypterous specimens previously have been known for species of Zetekella and Minitingis. We do not agree with the terminology typically used in the specialized literature to differentiate these two wing forms, but we reserve this subject for a more comprehensive, and illustrated, treatment in a future contribution. Froeschner (1968) noted that only Minitingis and Gonycentrum Bergroth have seven cephalic spines in Phatnomatini, assuming that Zetekella has only five. Drake (1944), however, in describing the genus and Z. zeteki, already had observed that "there are indications of a pair of spines on the head behind the eyes and just in front of the collar" and that "as these are very much atrophied, they are not mentioned in the generic description." Because the type specimen housed at the USNH is missing the head, this statement could not be verified. This feature, however, could be seen in the voucher specimen for the new record. Moreover, these spines were also observed in the new species. Yet, the mistake was perpetuated in the identification keys of Froeschner (1996). Froeschner (1968) also delimited and revalidated Minitingis on the basis of the acute processes of the paranota, which, however, can vary (Barber 1954).
In addition to cephalic spines and pronotal processes, Froeschner (1968) used rostrum length and presence of an abdominal groove as characters that validate the genus Minitingis. These characters were not possible to observe in the holotype (and single known specimen) of M. elsae due to the way the specimen is mounted, but they could  be seen in all specimens of M. minusculus studied. We agree with Froeschner (1968) in regarding these two characters as reliable for distinguishing Minitingis from Zetekella. Froeschner's (1968) comments on the zoogeographical significance of the distributional records of both genera remain relevant following our description of a new species of Zetekella and report of new distribution records for Z. pulla and Z. zeteki.
Therefore, we still consider Minitingis a valid genus, but we expanded the diagnosis of Zetekella to include the occipital cephalic spines and removed the acute processes on the paranota as a reliable character for delimiting Minitingis.