Corresponding author: Danny Shpeley (
Academic editor: T. Assmann
Based on small samples (exemplars) analyzed with morphological methods, including detailed descriptions and illustrations, this study treats primarily the
The taxonomic investigation reported herein, began with discovery of an undescribed Mexican species of
Although the six known Brazilian species had been clearly described and well illustrated, a more detailed comparison of
We broadened our study by placing
This study is based on examination of 94 specimens of
Measurements. Measurements were made with an ocular micrometer in a Wild M5 stereoscopic microscope, at 50×. Measurements of external body parts and abbreviations used for them in the text are:
Length of head - linear distance from apex of extended left mandible to posterior margin of the postocciput;
Width of head - maximum distance across head, including eyes;
Length of antennomere 1 - linear distance from base of antennomere 1 to apex of antennomere 1;
Length of antennomeres 2–4 - linear distance from base of antennomere 2 to apex of antennomere 4;
Length of pronotum - linear distance from anterior to posterior margin, measured along the midline;
Maximum width of pronotum - greatest linear transverse distance;
Length of elytra - linear distance from humerus to apex;
Width of elytra - maximum distance across the elytra.
Overall Body Length is the sum of HL, PL, and EL. Values for ratios for species were computed, using the measurements above: A1L/A2-4L; HW/PW. These numerical data are illustrative rather than definitive.
To express quantitatively proportions of the phallus, three measurements were made, using left lateral and dorsal aspects as illustrated in
Length of phallus - measured in a straight line from basal to apical margin;
Length of apical portion - measured in a straight line from apical margin of periostial area to apical margin;
Width of phallus - maximum transverse distance across the shaft, in ventral aspect.
These measurements were combined as two ratios PAL/PL; and PSW/PL. These numerical data are illustrative rather than definitive.
Preparation of material. Dissections were made by using standard techniques. Genitalia and other small structures were preserved in glycerine in microvials and pinned beneath the specimens from which they were removed. Larger structures and those that were gold-coated for study with the SEM were glued to cards pinned beneath the specimens from which they were removed.
Micrographs of isolated structures were taken with a JEOL JSM 6301 FXV field emission SEM. Line drawings of selected body parts were prepared by using a camera lucida on a Wild W5 stereoscopic microscope. Plates were prepared by using Adobe Photoshop CS 4.
Citation of figures. Figures included in the present publication are cited in the text as “Fig.” Those previously published are cited “fig.”
Label data. For type material, the information on each label is reproduced as exactly as is possible using ordinary type. Information on each label is enclosed in quotation marks; as well, a semicolon marks the end of a label. A slash mark (/) indicates the end of each line of text.
Most of the terms used to designate details of structures are found in textbooks of general entomology, or are used by coleopterists, generally. Other words, used to designate particular structures or parts thereof, are not in general use, though they have been used by us in previous publications. We provide information about these words here, as well as names that have been changed for certain structural features.
Microsculpture. A “sculpticell” is the space on the surface of the cuticle enclosed by adjacent microlines of the integumental system of microsculpture (
Chaetotaxy. This term refers to the so-called “fixed setae”, which are the long, evidently tactile, commonly encountered setae on carabids: dorsal labral; clypeal; supraorbital; stipital; submental; mental; glossal; palpigeral; pronotal; elytral parascutellar, discal, and umbilicate (or lateral); coxal, trochanteral, femoral, and tarsomeral; abdominal sternal ambulatory (sterna IV, V, VI); and abdominal sternal terminal (sternum VII, near posterior margin). Standard leg setae were not included because of difficulty in distinguishing them from the general body setation; but see
Body parts. The term “segment” is restricted for use to those body parts that reflect embryonic somites; thus, somite-like portions of the abdomen are referred to as segments. Abdominal segments are designated by Roman numerals corresponding to their respective somites. The first complete sternum is III, and the last one normally exposed is VII. For numbering the genital somites, we follow
Head. The term “head capsule” in the
Eyes. Three conditions are recognized in the
Antennae. Antennomere 1 (
Mandibles.
Labium. The labium of the
Male tarsal vestiture. Two types of adhesive vestiture on the ventral surface of the fore tarsi are exhibited by males of the
Male genitalia. Based on economy of expression, in preference to the widely used terms “median lobe” and “internal sac”, we use “phallus” and “endophallus”, respectively. Such usage is also well established in the entomological literature treating the male reproductive system. The surface of the phallus treated by convention as dorsal is really the ventral surface, and vice versa (
Ovipositor (
Some of the morphological features that distinguish
Comparison of diagnostic character state combinations exhibited by exemplars of
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01 | Antennomere 1, row of erect setae | character state 01.1<br/> absent | character state 01.2<br/> present<br/> |
character state 01.2<br/> present<br/> |
02 | Posterior supraorbital seta | character state 02.1<br/> absent | character state 02.2<br/> present<br/> |
character state 02.2<br/> present<br/> |
03 | Prosternal setae | character state 03.1<br/> absent | character state 03.2<br/> present | character state 03.2<br/> present<br/> |
04 | Male adhesive setae | character state 04.1 biseriate | character state 04.2<br/> pad-like,<br/> |
character state 04.2<br/> pad-like |
05 | Male genitalia, paraostial sclerites | character state 05.1<br/> present<br/> |
character state 05.2<br/> absent | character state 05.2<br/> absent |
06 | Female genital tract, bursal sac | character state 06.1<br/> absent | character state 06.2<br/> present<br/> |
character state 06.2<br/> present<br/> |
07 | Female genital tract, sec. spermathecal gland | character state 07.1<br/> present<br/> |
character state 07.2<br/> absent | character state 07.2<br/> absent |
08 | Integument color | character state 08.1<br/> rufous to piceous | character state 08.1<br/> rufous | character state 08.2<br/> testaceous |
09 | Head capsule form | character state 09.1 trapezoidal | character state 09.1 trapezoidal<br/> |
character state 09.2<br/> oviform<br/> |
10 | Eyes | character state 10.1 macrophthalmous | character state 10.1 macrophthalmous<br/> |
character state 10.2 microphthalmous<br/> |
11 | Metepisternum | character state 11.1<br/> elongate | character state 11.1<br/> elongate | character state 11.2<br/> quadrate |
12 | Hind wings | character state 12.1 macropterous | character state 12.1 macropterous | character state 12.2 brachypterous-apterous |
13 | Ovipositor: gonocoxite 2 | character state 13.1<br/> distal margin evenly rounded | character state 13.1<br/> distal margin evenly rounded<br/> |
character state 13.2<br/> distal margin deeply notched<br/> |
Included are two precinctive genera of
Habitus, dorsal aspect, of:
Head, dorsal aspect, of:
Head and prothorax, lateral aspect, of:
SEM micrographs of head and antennomere 1, dorsal aspect.
SEM micrographs of labra.
SEM micrographs of mandibles of
SEM micrographs of mandibles of
SEM micrographs of left maxillae and labia.
Digital images of male genitalia.
SEM micrographs of female ovipositor sclerites.
Line drawings of female reproductive tract, ventral aspect.
Digital images of male genitalia and line drawing of female reproductive tract of
Outline map of southern North America, Middle America and northern South America, showing generalized geographical range of species of
List of names of species of
As explained by
With character states of Western Hemisphere
None required here. See
Confined to the Neotropical Region, the seven species of this genus are known only from southeastern Brazil (
The previously described species of genus
Some specimens of
The ultrastructural features (i.e., principally sensillar) observed with scanning electron microscopy by
In contrast, the standard structural troglobitic features of
The external features of
Discovery of the Mexican species adds important details to the story of evolution of
An additional observation relating to evolutionary history of the
1 | Anophthalmous ( |
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1’ | Microphthalmous. Maximum width of elytra near middle, or posteriad middle. Male genitalia: right paramere styliform or not, distinctly shorter than left paramere | 2 |
2(1’) | Elytron with apical margin truncate, not sinuate. Male right paramere styliform or broad | 3 |
2’ | Elytron with apical margin sinuate. Male right paramere broad, not styliform, distinctly shorter than left paramere | 5 |
3(2) | Head dorsally without setae posteriad the anterior supraorbital setae (Pellegrini and Ferreira 2011: 49, fig. 2A) | |
3’ | Head dorsally with one to three pairs of setae posteriad the anterior supraorbital setae | 4 |
4(3) | Labrum with anterior margin broadly concave. Prosternal setae two pair ( |
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4’ | Labrum ( |
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5(2’) | Head dorsally with three pairs of setae posteriad the anterior supraorbital setae ( |
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5’ | Head dorsally with one or two pairs of setae posteriad the anterior supraorbital setae | 6 |
6(5’) | Head dorsally with two pairs of setae (posterior supraorbitals and occipitals,) at posterior border of head capsule ( |
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6’ | Single pair of setae (posterior supraorbitals) at posterior border of head capsule ( |
Only the following new species is treated here.
Three specimens, as follows. HOLOTYPE female, labeled: “MEXICO, Oaxaca/ 7100’ 21.8 mi./ n. Juchatengo/ VII.18-19.1966”; “George E. Ball/ D. R. Whitehead/ collectors” (USNM). PARATYPES two: female, labeled same as holotype (UASM); male, labeled: “MEXICO, Oaxaca/ Mt. Alban, near ruins/ Acacia scrub 6000 ft/ VI-3/4-82/ Rolf L. Aalbu, col.” (USNM).
Ridge top, Sierra de Miahuatlan, in western Oaxaca, Mexico, at 2164 m, 35 km north of San Pedro Juchatengo,
A Latinized eponym, masculine gender, genitive case, based on the surname of Donald R. Whitehead, now deceased, one of the collectors of the type series of this species.
See key, above.
Rectangular, lateral margins rounded; anterior margin irregularly convex; six dorsal setae, lateral setae longer than four medials. Epipharynx (
The specimens from the type locality were collected in the remains of a cloud forest. One was on the ground, in leaf litter; using a Bowie knife, the other was dug out of wet wood of a pine log, near the ground surface. The Monte Alban specimen was likely taken from a dark crevice, “where the temperature was much cooler than outside“ (Rolf Aalbu, personal communication). Judging from the structural features of the type material, this species may be troglophilic, but not troglobitic, as are its Brazilian congeners (
Attached to the holotype was a fungus of the species
(
Type specimens, only. For label details, see above.
A compound Latinized noun, treated as neuter, from the generic name
With character states of Western Hemisphere
None required here. See description of
Members of this genus are mesophilous to hygrophilous, occupying wet meadows and flood plains principally in open sites, but also in shaded areas along streams, in tropical gallery forest. For more details, see
For
Indicated by
A Latinized eponym, nominative case, based on the name of the country in which the type locality is located.
This species ranges from northern Mexico (states of Nuevo Leon, Nayarit, and Tamaulipas) northward to southwestern U.S.A (states of Arizona, New Mexico, and Texas;
Treatment of the Western Hemisphere zuphiines remains incomplete. The large geographical gap between the Brazilian and Mexican species of
At hemisphere level,
The authors are pleased to thank: Rolf L. Aalbu (Entomology, California Academy of Sciences, San Francisco California) for information about the locality where he collected a paratype of
Based on morphological features, most authors have treated the
Specimens of this tribe are small in size (overall length less than 7.00 mm), integument various (nearly white, testaceous to piceous), and generally pilose, head more or less constricted posteriorly, antennomere 1 (scape) elongate more than length of antennomeres 2+3 and wider than antennomeres 2-11; elytra with apical margin truncate, or subtruncate-sinuate, with a narrow membranous fringe; elytral interval 3 without fixed setae; tibiae without prominent spines. For a more detailed characterization of
The range of this tribe in the Western Hemisphere is co-extensive with the range of the
Based on previous studies,
The following key distinguishes among the Western Hemisphere genus-groups and genera of
1 | Maxillary and labial palpi similar to one another in size and proportions; maxillary palpomere 4 ( |
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1’ | Maxillary palpi much larger than labial palpi; maxillary palpomere 4 markedly enlarged. Head posteriorly less constricted. Male fore-tarsomeres 1-3 with adhesive vestiture biseriate, or a single row of squamo-setae | 3 |
2(1’) | Eyes absent or flat, ommatidia not evident. Metepisternum about quadrate. Elytra with humeri markedly constricted, lateral margins broadly rounded ( |
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2’ | Eyes convex, normal size, ommatidia evident. Metepisternum elongate, longer than wide at base. Elytra with humeri broadly rounded, lateral margins straight, parallel to one another ( |
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3(2’) | Head posteriorly constricted as very narrow neck (more or less as wide as diameter of one eye (cf. |
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3’ | Head posteriorly with neck thick (wider than diameter of one eye). Gonocoxite 2 with broad ensiform setae ( |
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4(3) | Elytron with lateral margin straight for most of length. Pronotal posteriolateral angles long, very prominent spines. Labium with submentum and mentum fused ( |
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4’ | Elytron with lateral margin broadly rounded. Pronotal posteriolateral angles short spines. Labium with submentum and mentum separated by a suture | |
5(3’) | Pronotal posteriolateral angles not spined | |
5’ | Pronotal posteriolateral angles each a small, sharp spine | 6 |
6(5’) | Antennal socket limited above and below by a sharp carina, inferior carina better developed than superior one (clearly visible from above). Antennomere 1 relatively shorter (slightly shorter than antennomeres 2-4). Pronotum as long as wide; anterior angles more or less sharp. Integument generally glabrous | |
6’ | Antennal socket with superior and inferior carinae equally developed. Antennomere 1 relatively longer (as long or longer than antennomeres 2-4). Pronotum longer than wide. Pronotum and elytra setose |