Corresponding author: Emily A. Hartop (
Academic editor: Torsten Dikow
A new
The genus
The following represents a newly discovered species with wing spots that make it an easily recognized species of
Specimens were collected from the Zurquí All
In work on the ZADBI Project and beyond, the authors have spent countless hours sorting through tens of thousands of worldwide
The task of developing an unfailing system of description for organisms as diverse and numerous as phorids, is a problem opined as impossible by Malloch (
Specimens were collected from Zurquí de Moravia, San José, Costa Rica in Malaise Traps, preserved in 95% ethanol, and subsequently HMDS dried (
Standard morphological terminology, as found in the Manual of Central American
Common terms like “hairs” and “bristles” are used to delineate different types of socketed setae. “Hairs” we use to refer to smaller, non-feathered setae, while “bristles” are much larger, more robust and feathered setae. Although both categories can range in length and thickness, the feathering on a true “bristle” can be readily observed on a standard compound microscope at 40×.
The presented character set and states were developed by combining the classic characters outlined by those such as Lundbeck (
The authors had found certain characters useful to organize large groups of
As this system had proven utility in their own studies, the authors decided to formally present a new format for published descriptions of this fauna based on ease of use, and in anticipation of the tremendous number of descriptions forthcoming in this genus. The system of description that is proposed organizes key characters into tables supplemented by visual representations of genitalia and any other salient features, as well as clear habitus, hypandrium and wing photographs. While the authors do not intend the tables themselves to be definitively diagnostic in all cases, the combination of the table and visual aids will be so, and will replace unnecessarily verbose descriptions. This will allow any user to efficiently not only compose new descriptions for the fauna, but to more easily scan vast amounts of existing literature composed in this manner.
In addition to tabulated descriptions, all character states have been coded for use in a character state matrix that will be searchable as an online guide (ie. character "x" has states "a", "b", etc.). This online system (found at
Rare
Lastly, characters that are definable but not practically usable (such as absolute colors, which would require the use of color palettes accounting for qualities such as hue, tint, shade, and tone while accommodating variation in digital viewing, printing, and specimen lighting), are omitted. Relative coloration (such as halter to scutum) is much less subjective and easily used for coding. It is important to note that relative coloration will occasionally create a discrepancy between a coded character state and what a user observes. Examples of dark species with dark halters that code as “lighter”, because they happen to be lighter than the scutum, do occur. Similarly, light species with light halters that happen to be slightly darker than the scutum and therefore key as “darker” do exist. The authors therefore include the relative coloration of halter to scutum as an easy to use, unambiguous character that agrees with visual perception in most cases. The habitus photographs included with each description will serve as an excellent way to easily compare qualitative characters, including color, for many parts of the specimen.
In order to utilize this description method, specimens should ideally be available in chemically dried, slide mounted, and ethanol preserved (for dissection) states. Habitus photos of chemically dried specimens are essential for conveying the overall coloration, shape, and gestalt of a species while more detailed characters must be observed on slide mounted specimens with a compound microscope. A photograph of the dissected hypandrium can be an essential tool for identification, and this requires additional specimens available for dissection. Therefore, with this system the ideal number of specimens would be a minimum of three. It is the practice of the authors in cases where limited specimens are available to photograph the specimens for habitus images while they are still in ethanol and then slide mount those specimens for detailed observations and measurements. On occasions where only a single specimen is available, this may mean the omission of a separate, dorsoventrally oriented hypandrial photograph.
Template for
Descriptions of characters/states:
Supra-antennal (SA) setae ratio: Rather than use the more subjective visual comparisons used classically, the length of the ventral SA are entered as a percentage (given as a decimal to second digit) of the length of the dorsal SA. Therefore, SA that are equal in length will have a value of “1.00”, ventral SA that are half the length of the dorsal SA will have a value of “0.50”, etc.
Positioning of ventral interfrontal (VIF) setae: Coded as either ”Normal” orientation (a) or “VFO adjacent” (b). Normal orientation is indicated by the ventral interfrontal (VIF) setae being located either roughly midway between the supra-antennal (SA) setae and the ventral fronto-orbitals (VFO), or closer to the SA (Fig.
Postpedicel Subcuticular Pit Sensillae (SPS) vesicles: absent (a) or present (b) (
Palpal setae: “long” (a) if longer than the width of the palpi, “short” (b) if shorter than the width of the palpi.
Labellum: “not spinose” (a) if spinose setulae not present, “sparse” (b) if setulae are scattered and present in numbers fewer than 30 per labellum, “dense” (c) if setulae are densely covering labellum (
Anepisternum: An anepisternum without setae is scored as “bare” (a). If setae are present, they can be “hairs only” (b) or they can be “hairs + bristle(s)” (c). If bristles are present, the number and size may be indicated (ie. “hairs + one long bristle”) as a remark, that data will not be coded.
Halter color: halter color is based on the knob of the halter when compared with the color of the scutum. Halteres that are lighter than the scutum are coded as “lighter” (a), halteres that are the same color as the scutum key as "same", and halteres that are darker than the scutum are coded as “darker” (c). It is important to note that in light colored species, halteres may be quite light in color in an absolute sense, but in a relative sense will be scored in the “darker” category, or vice versa. Halter color is best observed on either dried specimens or specimens preserved in ethanol.
Number of notopleural (NP) setae: scored as 2 (a) or 3 (b).
Notopleural (NP) cleft: scored as absent (a) or present (b).
Scutellar setae: scored as “4 =” (a) if anterior and posterior are the same size, “4 /=” [note "/=" is a more computer-friendly version of the mathematical "not equal" symbol], (b) if the anterior and posterior pairs are not the same but the smaller pair are longer than the posterior small setulae on the scutum, and “2+2” (c) if the smaller pair are minute and equal in length to the posterior small setulae on the scutum.
Foretarsus (ts1) palisade: indicate the segment numbers of the foretarsus on which a setal palisade is present (i.e. Tarsomeres 1–4, etc.).
Midtibial (t2) palisade: indicate portion of tibia on which a setal palisade is present (0.50, 0.75, etc.).
Bifurcated spines in comb of hind tibia (t3): “absent” (a), or “present” (b).
Hind tibial (t3) setulae: a single, posterodorsal, row of setulae on the hind tibia is “PD only” (a); if a second row is present on the anteroventral side of the tibial palisade, that is “PD + AD” (b).
Hind femoral (f3) basal setae: length of basal setae on hind femur indicated relative to setae on the anteroventral surface of the hind femur as “B<AV” (a), “B=AV” (b) or “B>AV” (c).
Hind femoral (f3) basal setae differentiation: “absent” (a) or “present” (b), description in remarks. Examples of differentiation would be setae that are clearly thickened, curved, found in unusual configurations, etc.
Wing length: given in mm to second decimal, measured as per Schmitz (fig. 45 in
Subcosta (Sc): given as “complete” (a) when it reaches vein R1 or “incomplete” (b) when it fades out before reaching R1.
Hair at base of vein R: either “absent” (a), “minute” (b) if shorter than the width of the R vein, “short” (c) if longer than the width of the vein but shorter than 2× the width of the vein, or “long” (d) if longer than 2× the width of the vein.
R2+3 vein: either “present” (a) or “absent” (b).
Costal index (CI): given as a decimal to second digit, found by dividing the wing length by the length of the costal vein (fig. 45 in
Costal ratios: given as “C1:C2:C3” with C3 at a value of 1 and others to second decimal (fig. 44 in
Costal setae length: measurement in mm of the length of the longest costal setae present in section 3 of the costa.
Number of alular setae: given as a number, coded as “1” (a), “2” (b), “3” (c), “4+” (d) due to observed variation in number of alular setae on species with more than four present.
Alular setae length: given in mm, longest seta present is measured.
Wing color: either “lightly infuscated/clear” (a) or “strongly infuscated” (b); the authors understand that this character may include some ambiguity, so therefore it must be clarified that most wings are state "a", and only wings that are distinctly infuscated should be coded as the second character state. Wings are best viewed on dried specimens (or in a clear mounting medium, ie.
Anal tube (AT) length relative to length of dorsal face of epandrium (E): length of the anal tube (cerci + hypoproct) is given relative to the dorsal face of the epandrium as “AT<E” (a), “AT=E“ (b), “AT>E” (c). This comparison is most easily done on a slide mounted specimen, where the epandrium can be seen even if it is partially covered by the tergites and must be viewed through them.
Epandrial (E) Setation: “hairs only” (a) or “hairs + bristle(s)” (b).
Relative lengths of posterior setation: indicate the relative lengths (not thickness) of setae found on Tergite 6 (T6), Epandrium (E), Cerci (C) and Hypoproct (H) with operators < [less than], = [equal], or ~ [subequal: approximately equal] (ex. T6<C=H<E).
See Table
Wing with darkly-pigmented central swelling in center. Fork formed by wing veins R2+3 and R4+5 greatly enlarged. The central wing spot makes this different from all other described Neotropical species with the single exception of
Named for E. A. Hartop's niece, Shade Zehendner.
Known from a single site in Costa Rica.
Unknown.
A primary key to Neotropical species of
Neotropical species of
Three genera that have been synonymized (or partially synonymized) with
The presence of such a remarkable and distinct undescribed species of Neotropical
We are grateful to the staff of the ZADBI project for their hard work and professionalism: Carolina Avila, Estella Hernandez, Anna Holden, Marco Moraga, Annia Picado, Wendy Porras, Elena Ulate, and Elvia Zumbado. We thank Lisa Gonzalez for her technical assistance. R.H.L. Disney is thanked for his comments on an early version of this manuscript, as are reviewers of the manuscript whose comments helped to improve upon it greatly. Funding was provided by US National Science Foundation grant DEB-1145890 to B. Brown and A. Borkent. This is contribution No. 3 of the ZADBI project.
Frontal setation patterns showing "normal" (a & b) and "VFO adjacent" (c & d) arrangements. vs = vertical seta, po = postocellar seta, dif = dorsal interfrontal seta, dfo = dorsal fronto-orbital seta, dsa = dorsal supra-antennal seta, vfo = ventral fronto=orbital seta, vif = ventral interfrontal seta, vsa = ventral supra-antennal seta.
Template for
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SA ratio | decimal to second digit | |
VIF position | normal or VFO adjacent | |
SPS vesicles | absent or present | |
Palpal setae length | long or short | |
Labellum spinosity | not, sparse, or spinose | |
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Anepisternum | bare, hairs only, or hairs + bristles | |
Relative halter color | lighter, same, or darker | |
# NP setae | number | |
NP cleft | absent or present | |
Scutellar setae | 4 =, 4 /=, or 2+2 | |
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ts1 palisade | number of tarsomeres | |
t2 palisade | portion of tibia present (decimal to second digit) | |
t3 comb bifurcate | absent or present | |
t3 setulae | PD or PD+AD | |
f3 basal setae | B <, =, or > AV | |
f3 basal setae differentiation | absent or present | |
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Fig X | |
Wing Length (mm) | decimal to second digit | |
Subcosta | complete or incomplete | |
Hair at base of R | absent, minute, short or long | |
R2+3 | present or absent | |
Costal index | decimal to second digit | |
Costal ratios | C1:C2:C3 (C3=1) | |
Costal setae length (mm) | decimal to second digit | |
Number alular setae | number | |
Alular setae length (mm) | decimal to second digit | |
Wing color | lightly infuscated/clear or strongly infuscated | |
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Fig X, X (genitalia, hypandrium) | |
AT length | AT <, =, or > E | |
E setation | hairs only or hairs + bristles | |
Relative posterior setation | T6, E, C, H in ascending order using (<, =, ~) | |
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any information unique to species |
Description of
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SA ratio | 1 | |
VIF position | VFO adjacent | |
SPS vesicles | absent | |
Palpal setae length | long | |
Labellum spinosity | spinose | |
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Anepisternum | bare | |
Relative halter color | same | |
# NP setae | 2 | |
NP cleft | absent | |
Scutellar setae | 2+2 | |
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ts1 palisade | 1-4 | |
t2 palisade | 0.67 | |
t3 comb bifurcate | absent | |
t3 setulae | PD | |
f3 basal setae | B<AV | |
f3 basal setae differentiation | absent | |
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Fig. |
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Wing Length (mm) | 1.56 | |
Subcosta | incomplete | |
Hair at base of R | long | |
R2+3 | present | large fork |
Costal index | 0.65 | |
Costal ratios | 1.25:1.00:1 | |
Costal setae length (mm) | 0.09 | |
Number alular setae | 3 | |
Alular setae length (mm) | 0.15 | |
Wing color | strongly infuscated | |
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Figs |
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AT length | AT>E | |
E setation | hairs only | |
Relative posterior setation | T6~E~H<C | |
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wing with central, bubbled, pigmented spot |