Western Hemisphere Zuphiini: descriptions of Coarazuphium whiteheadi, new species, and Zuphioides, new genus, and classification of the genera (Coleoptera, Carabidae)

Abstract Based on small samples (exemplars) analyzed with morphological methods, including detailed descriptions and illustrations, this study treats primarily the Zuphium genus-group in the Western Hemisphere, which comprises two precinctive genera: Coarazuphium Gnaspini, Vanin & Godoy, 1998 (type species Parazuphium tessai Godoy & Vanin, 1990) and Zuphioides gen. n. (type species Zuphium mexicanum Chaudoir, 1863). The genus Coarazuphium includes six troglobitic species from Brazilian caves, and one probably hypogaeic (troglophilic) species from the mountains of Oaxaca, in Mexico (Coarazuphium whiteheadi, sp. n., type locality, ridge top, in western Oaxaca, Mexico, at 2164 m, 35 km north of San Pedro Juchatengo, 16.462N, 97.010W). The epigaeic genus Zuphioides includes 23 species, with its geographical range extended from Neotropical temperate Argentina in southern South America, northward through the tropics to north temperate southeastern Canada, in the Nearctic Region. Keys are provided to the species of Coarazuphium and to thegenera of Western Hemisphere Zuphiini.


HL
Length of head -linear distance from apex of extended left mandible to posterior margin of the postocciput; HW Width of head -maximum distance across head, including eyes; A1L Length of antennomere 1 -linear distance from base of antennomere 1 to apex of antennomere 1; A2-4L Length of antennomeres 2-4 -linear distance from base of antennomere 2 to apex of antennomere 4; PL Length of pronotum -linear distance from anterior to posterior margin, measured along the midline; PWM Maximum width of pronotum -greatest linear transverse distance; EL Length of elytra -linear distance from humerus to apex; EW Width of elytra -maximum distance across the elytra.

OBL
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 Figs 10 and 13:

PL
Length of phallus -measured in a straight line from basal to apical margin; PAL Length of apical portion -measured in a straight line from apical margin of periostial area to apical margin; PSW 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 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.

Structural features
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 (Allen and Ball 1980: 485-486).In most groups, overall, microsculpture is varied.In contrast, in the Zuphium species group, it is absent (surface smooth) or simply isodiametric, with microlines very fine and sculpticells flat to slightly granulate.Emphasis is placed on description of microsculpture of the sclerites of the dorsal surface, which is adequate for characterization of the taxa of the Zuphium genus-group.
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 Bils (1976).
Head.The term "head capsule" in the Zuphium genus-group is restricted to that portion of the head that extends from the anterior margin of the clypeus to the broad part of the occiput that precedes the more or markedly narrowed posterior part of occiput + postocciput.
Mandibles.Shpeley and Ball (2001: 9-21, figs 3A-E) characterized the mandibles of the lebiine subtribe Pericalina, and illustrated the major features with SEM figures.We use the same system here.See Figs 6A-F and 7A-F.
Labium.The labium of the Zuphium genus-group is standard for Carabidae.For the combined glossae and paraglossae, we use the standard term ligula.The central sclerotized, apically setigerous, structure is the glossal sclerite.See Figs 8C and 8F.
Male tarsal vestiture.Two types of adhesive vestiture on the ventral surface of the fore tarsi are exhibited by males of the Zuphium genus-group: articulo-setae (Fig. 9A, as) and biseriate squamo-setae (See Stork 1980, for a thorough discussion of this topic).
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 (Deuve 1993: 88).As in our previous publications, we have chosen to remain with the conventional usage.Phalli were classified as anopic, with the ostium dorso-medial.The phallus (e.g., Figs 10A-10F, and 13A-13C), illustrated in left lateral, ventral and right lateral aspects, with base toward the bottom of the page, exhibits interspecific differences in form.These differences are seen readily as overall patterns ('Gestalt') but are not so easily described, except with notation of differences in size and shape of the apical area.To provide the basis for verbal description, two principal regions are distinguished--the shaft (s), and basal bulb (bb), or lobe, or phallobase.The latter is the swollen area set at an angle to the ventrally curved shaft, surrounding the basal opening; the periostial membrane (om), which surrounds the ostium, marks the place of egress of the endophallus during copulation; an apical portion (ap) or phalloapex, extends distally from the apex of the periostial membrane to the apex of the phallus.The phalli in lateral aspect exhibit curvature ventrally.A pair of slender paraostial sclerites (ps) is evident (Figs 13A,13C) or not (Fig. 10A).
Ovipositor (Figs 11A-11F) and female genital tract (Figs 12A,12B and 13D).For naming the sclerites of the adephagan ovipositor and units of the female genital tract, we follow the system used by Liebherr and Will (1998;see figures therein).For the gonocoxites (gc1, gc2), the surfaces that are ventral in the infolded position are lateral when the ovipositor is extended; thus such surfaces are designated as lateral, and the other surfaces are designated accordingly.systematics Some of the morphological features that distinguish Coarazuphium from the Eastern Hemisphere widespread Z. olens species group (includes Z. olens (Rossi, 1790), type species of the genus) are shared with the Western Hemisphere species of Zuphium of authors (Table 1).From that distribution of character states, we conclude that the Western Hemisphere Zuphium of authors forms an assemblage different from Coarazuphium and from the Z. olens species group, and this bridges the morphological gap between those two taxa.A reasonable taxonomic expression of this situation seems to us is to recognize three groups of generic rank: Zuphium (sensu stricto), Zuphioides new genus (= Zuphium of auothors), and Coarazuphium.Microsculpture.Dorsal surface of head (including clypeus) and pronotum smooth, microlines not evident; labrum and elytra with microlines fine (not easily seen at 50× or lower magnification), mesh pattern isodiametric.Ventral surface with microlines fine, generally transverse, sculpticells rather short.
Luster.Dorsal surface of head and pronotum shiny, elytra somewhat duller.
Geographical distribution.The range of this genus-group in the Western Hemisphere extends from Neotropical Argentina (Reichardt 1977: 449) northward through the American tropics and subtropics to temperate eastern and central U.S.A., and to southeastern Canada (Lindroth 1969(Lindroth : 1090)), with an isolated population of Z. americanum in western Oregon (Bousquet, 2012(Bousquet, : 1356)).
Included taxa.Type species.Parazuphium tessai Godoy & Vanin, 1990(designated by Gnaspini et al. 1998).Generic name.As explained by Gnaspini et al. (1998: 299), the name Coarazuphium is a compound Latinized neuter noun derived from "coara", meaning hole or cave in the Tupi language (Brazilian, native tongue), plus Zuphium.The name refers to the troglophilic or troglobitic habits of the species of this genus, and their basic nominotypical Zuphium-like features.
Geographical distribution.Confined to the Neotropical Region, the seven species of this genus are known only from southeastern Brazil (Pellegrini and Ferreria 2011b: 57, fig. 10) and southern Mexico (Map.1).This is an example of the "Paleo-American distribution pattern" (see Halffter (1987) and Liebherr (1994: 845)).
Way of life.The previously described species of genus Coarazuphium inhabit caves in one of two types of substrate (Pellegrini and Ferreira 2011b:  The caves in iron ore are described as "shallow".Members of those species occupying the caves in limestone substrate are described as freely walking over the soil, and presumably resting exposed, rather than resting (hiding?) under rock cover.In contrast, all of the specimens collected of C. tapiaguassu were found under rocks, on the cave floor.In contrast, known members of the Mexican species, C. whiteheadi, new species, seem to be surface inhabitants (see below) Parasites.Some specimens of C. tapiaguassu were infested by laboulbenelian fungi.The fungi were not identified further.See this topic for C. whiteheadi, below.
Evolutionary considerations.The ultrastructural features (i.e., principally sensillar) observed with scanning electron microscopy by Pellegrini andFerreria (2011a, 2011b) differ only slightly between the two species of Coarazuphium that they studied, and between that genus and Zuphioides (cf.accompanying SEM illustrations of Z. mexicanum).So, it seems to us unlikely that such features are or will be evolutionarily informative.
In contrast, the standard structural troglobitic features of Coarazuphium (lengthening of antennae and legs, depigmentation, micro-or anophthalmy, and reduction of elytral length, elytral humeri, metathorax and metathoracic wings) plus details of male and female genitalia are evolutionarily informative.Based on eye loss and more elongate appendages, Gnaspini et al. (1998: 303) proposed that C. cessaima showed the more modified features, compared to C. tessai and C. bezerra, the only other species of Coarazuphium known at that time.Álvares and Ferreira (2002: 43) proposed that, based on the features noted above, their newly described C. pains would occupy a position intermediate between C. cessaima and C. tessai + C. bezerra.The external features of Coarazuphium evidently evolved in parallel with, and independent of, four other zuphiine troglobite taxa: the remarkable Spanish Ildobates neboti Español, 1966; the Canary Islands Parazuphium feloi Machado (1997: 163); the Austral-ian Nullarbor Speozuphium poulteri Moore (1995: 159) and Speothalpius grayi Moore (1995: 160).Another Canary Islands Parazuphium (P.damascenum canariense Machado, 1992: 580) exhibits these same reductive (though less developed) features, but it is evidently conspecific with the continental Palaearctic P. damascenum damascenum (Fairmaire, 1897).For a more detailed discussion of the matter see Machado (1992: 581-582).Gnaspini et al. (1998: 308-309) proposed for the Brazilian species of Coarazuphium that "..... these highly derived troglobitic features are due to a long-term isolation   inside the subterranean environments, which took place under the drier climate to which the region was in the past and is still submitted in the present.It is largely accepted in the literature that cave arthropods are related to litter epigean and/or endogean ancestors, which already inhabited humid habitats.Therefore, ...... the ancestral species [of Coarazuphium] should have been epigean and lived in forested (or at least humid) areas, and occurred at least in part of the region where the genus occurs nowadays.From [such ancestral stock] several lineages invaded the caves from the northern Bambui Speleological Province, where they became isolated with the progressive shrinkage of humid environments.Thence, the origin of the genus takes back to a time when the area was not drying yet, which is probably the Tertiary".
Discovery of the Mexican species adds important details to the story of evolution of Coarazuphium.First, the marked geographical range extension (from southeastern South America to the southern part of the North American continent) shows that this genus was not confined to eastern Brazil.Further, the distribution pattern lends support to the hypothesis of Gnaspini et.al. (see above) that Coarazuphium originated in early Tertiary time (i.e., before the beginning of the drying trend that extended through much of the Cenozoic era).Second, the extra-speleal humid forest existence of C. whiteheadi suggests that the basic troglobitic features of Coarazuphium (microphthalmy, brachyptery, depigmentation, etc.) evolved in surface habitats, though probably in forested deep leaf litter locations, and were in effect preadaptations for cave life.
An additional observation relating to evolutionary history of the Zuphium genus group is that shared features of Zuphioides and Coarazuphium indicate that these two genera may be adelphotaxa, with Zuphioides retaining mostly ancestral features including life in lowland hygrophilous or mesophilous situations.
Microsculpture.Dorsal surface of head capsule (including clypeus) and pronotum smooth, microlines not evident; labrum and elytra with microlines fine (not easily seen at 50× or lower magnification), mesh pattern isodiametric.Ventral surface with microlines fine, generally transverse, sculpticells rather short.
Collecting notes and habitat.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 (Gnaspini et al. 1998;Pellegrini and Ferreira 2011b).
Parasites.Attached to the holotype was a fungus of the species Rhachomyces zuphii Thaxter (Laboulbeniales: Laboulbeniaceae) (determiner of fungus not indicated on label), which is now attached to a pinned rectangular piece of clear plastic by a drop of balsam.
Geographical distribution (Map 1).This species is known only from two montane localities in the Mexican state of Oaxaca.
Description.None required here.See description of Z. mexicanum, below.

Zuphium mexicanum
Type area.Indicated by Mateu (1985: 330) as Veracruz, for Z. mexicanum; for Z. vicinum, "Mexico".Specific epithet.A Latinized eponym, nominative case, based on the name of the country in which the type locality is located.
Microsculpture.Dorsal surface of head capsule (including clypeus) and pronotum smooth, microlines not evident; labrum and elytra with microlines fine (not easily seen at 50× or lower magnification), mesh pattern isodiametric.Ventral surface with microlines fine, generally transverse.
Luster.Dorsal surface of head and pronotum shiny, elytra somewhat duller.
Elytra (Fig. 1B).Separate from one another, not fused along suture.Each elytron more or less rectangular, lateral margin straight; humerus projected anteriorly, basal ridge straight; apical margin truncate, with narrow band of membrane.Surface with striae very shallow, intervals almost flat.
Abdomen.As described for Western Hemisphere Zuphium genus-group.
Geographical distribution.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; Bousquet (2012Bousquet ( : 1357)).

Concluding remarks
Treatment of the Western Hemisphere zuphiines remains incomplete.The large geographical gap between the Brazilian and Mexican species of Coarazuphium causes one to wonder just how large the gap is.Surely, considering that this genus is likely to be an old (early Tertiary) resident of the Middle American highlands, we can expect therein additional species.
At hemisphere level, Zuphioides must be reviewed in detail, using as the principal basis the work of Joaquin Mateu (1981).No doubt, many more species remain to be discovered and described.From a World perspective, a generic-subtribal revision of the Zuphiini is required, based primarily on structural details of the male and female reproductive organs.
nian Institution, Washington, D. C.) for loan of specimens, and general advice about organization and structure of the manuscript; and David R. Maddison (Zoology, Oregon State University, Corvallis, Oregon), for advice about interpretation of molecular data in relation to classification of the Zuphiitae.Careful reviews of a previous draft of the submitted manuscript were made by our colleagues Douglas A. Craig and Bruce S. Heming.We are grateful for their considerable efforts, which led to a substantially improved manuscript.We are also grateful for the advice received from the referees, who reviewed the penultimate draft of our manuscript.
CS 4. Citation of figures.Figures included in the present publication are cited in the text as "Fig."Those previously published are cited "fig."

table 2 .
List of names of species of Coarazuphium Gnaspini, Vanin & Godoy and Zuphioides, new genus.