Parazuphium tessai Godoy & Vanin, 1990 (designated by Gnaspini et al. 1998).

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.

With character states of Western Hemisphere Zuphium genus-group, restrictedas follows: size small; pronotum and dorsal surface of elytra moderately densely setose, setae decumbent; body markedly depressed; integument pale; head narrow, oviform, with posteriolateral margins broadly rounded; antennae elongate, antennomere 1 as long or longer than antennomeres 2-4; humeri narrowed, slightly or markedly so; metasternum short, metepisternum quadrate. Brachypterous or apterous. Male genitalia (Figs 10A–10C): phallus without dorsal paraostial sclerites. Female genital tract (Fig. 12A, 12B): without secondary spermathecal gland.

None required here. See Gnaspini et al. (1998: 298-299), Pellegrini and Ferreira (2011b), and description of Coarazuphium whiteheadi below.

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)).

The previously described species of genus Coarazuphium inhabit caves in one of two types of substrate (Pellegrini and Ferreira 2011b: 55): limestone, occupied by Coarazuphium bezerra Gnaspini, Vanin & Godoy; Coarazuphium cessaima Gnaspini, Vanin & Godoy; Coarazuphium formoso Pellegrini & Ferreira; Coarazuphium pains Álvares & Ferreira; and Coarazuphium tessai (Godoy & Vanin); and iron ore, occupied by Coarazuphium tapiaguassu Pellegrini & Ferreira. 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 Coarazuphium tapiaguassu were found under rocks, on the cave floor. In contrast, known members of the Mexican species, Coarazuphium whiteheadi, new species, seem to be surface inhabitants (see below)

Some specimens of Coarazuphium tapiaguassu were infested by laboulbenelian fungi. The fungi were not identified further. See this topic for Coarazuphium whiteheadi, below.

The ultrastructural features (i.e., principally sensillar) observed with scanning electron microscopy by Pellegrini and Ferreria (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 Zuphioides 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 Coarazuphium cessaima showed the more modified features, compared to Coarazuphium tessai and Coarazuphium 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 Coarazuphium pains would occupy a position intermediate between Coarazuphium cessaima and Coarazuphium tessai + Coarazuphium 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 Australian Nullarbor Speozuphium poulteri Moore (1995: 159) and Speothalpius grayi Moore (1995: 160). Another Canary Islands Parazuphium (Parazuphium damascenum canariense Machado, 1992: 580) exhibits these same reductive (though less developed) features, but it is evidently conspecific with the continental Palaearctic Parazuphium 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 Coarazuphium 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 groupis 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.

Key to species of genus Coarazuphium Gnaspini, Vanin & Godoy

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, 16.462N, 97.010W.

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.

Size and proportions. Small, OBL 4.13–4.40 mm; EW 1.36–1.48 mm A1L/A2-4L 1.03–1.11; HW/PW 1.08.

Color. Body testaceous, in life almost white; appendages slightly paler; vestiture golden.

Habitus (Fig. 1A). Flat, overall. Head capsule (dorsal aspect, Fig. 2A) oviform, broadly rounded posteriolaterally. Eyes (Fig. 3A) flat, hardly perceptible, ommatidia not evident at 50×. Pronotum narrow. Elytra narrowed anteriorly and posteriorly, lateral margins distinctly bowed.

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.

Luster. Dorsal surface of head and pronotum shiny, elytra somewhat duller.

Body vestiture and punctation. Dorsal surface of head and pronotum sparsely punctate, vestiture sparse. Elytra with punctation and vestiture dense, vestitural setae decumbent. Abdomen. Abdominal sterna IV-VI with punctation rather sparse, vestitural setae decumbent.

Fixed setae. Head (Fig. 4A, 4C): clypeus one pair; head capsule with anterior pair of supraorbitals (asos) above eyes; posterior pair of supraorbitals (psos) posteriad eyes; one pair of postocular setae (pos) immediately behind eyes, laterally; one pair of occipital setae (ocs) posteriorly and mediad eyes; also one pair posterior supernumerary setae (psus) laterad and close to posterior supraorbital setae. Antennomere 1 (Fig. 4B, as1) with single long seta distally, and row of several semi-erect setae more proximally (Fig. 4B, as2, as3). Pronotum with two pairs of lateral marginal setae, anterior pair in anterior 1/8, posterior pair at posteriolateral angles. Prosternum with one pair paramedial setae anterioventrally (cf. Fig. 9B). Elytra: each elytron anteriorly with parascutellar seta; lateral setae about 21 (one group anteriorly, one group posteriorly, and single seta medially). Abdominal sterna IV–VI each with one pair of ambulatory setae, sternum VII with one pair of long setae near posterior margin.

Head, dorsal aspect (Fig. 1A). Postoccipital suture evident. Frontoclypeal suture present, with frons and clypeus separate. Genal sulcus (Fig. 4A, gs) broad, ventral margin sinuous.

Eyes. (Fig. 3A) Small, flat, ommatidia not evident at 50×, or lower magnification.

Antennae. (Fig. 4A). Filiform, extended about body length. Antennomere 1 (a1) rather slender, slightly longer than antennomeres a2-a4; antennomere 2 short, about half length of a3; antennomeres 3–11 narrow, cylindrical, distinctly longer than wide.

Mouthparts. As described for Zuphium genus-group, above. Labrum (Fig. 5A).

Rectangular, lateral margins rounded; anterior margin irregularly convex; six dorsal setae, lateral setae longer than four medials. Epipharynx (Fig. 5B).

Mandibles (Figs 6A–6F) Maxillae (Figs 8A–8B). Labium, ventral aspect (Fig. 8C).

Pronotum (Fig. 1A). Anterior margin truncate, lateral margins markedly sinuate posteriorly, posteriolateral angles prominent, dentiform, slightly anteriad posterior margin; surface impressions (anterior and posterior transverse and median longitudinal) shallow; lateral grooves and posteriolateral impressions moderately deep.

Pterothorax. Metasternum short; metepisternum quadrate.

Elytra (Fig. 1A). Separate, not fused along suture. Each elytron more or less rectangular, but narrowed anteriorly and posteriorly, lateral margin thus distinctly bowed; humerus projected anteriorly, basal ridge sinuate; apical margin truncate, with narrow band of membrane; surface with striae very shallow, intervals almost flat.

Hind wings. Short stubs, brachypterous.

Abdomen. Sternum VII with apical margin truncate.

Male genitalia (Figs 10A–10C). Anopic. Phallus narrow (PW/PL 0.238), slightly curved ventrally, narrowed apically, apical margin rounded, apical portion very short (AL/PL 0.063), ostial membrane extensive (OM/PL 0.317). Left paramere (lp) conchoid. Right paramere (rp) styliform, relatively long (RP/LP 0.317).

Female genitalia: ovipositor (Figs 11A–11C). Gonocoxite 1 (gc1) with patch of long trichoid setae distally on ventral surface. Gonocoxite 2 (gc2) short, thick; in lateral aspect falciform, apex pointed, row of four long trichoid setae dorso- and ventro-laterally; in dorso-ventral aspect, broad, paddle-like, apex deeply notched; dorsal surface glabrous except for dorso-lateral and ventro-lateral trichoid setae; ventral surface toward margins with row of pit pegs (mpp), preapical setose organ (pso) circuloid, with two furrow pegs (fp) and two very short nematiform setae (ns).

Female genital tract (Fig. 12A). Bursa copulatrix (bc) ended in an expanded bulbous anterior extension (bs). Common oviduct (co) inserted in bursa copulatrix at base of the anterior extension. Spermatheca (sp, broken) slender, inserted on bursal sac beside insertion point of spermathecal gland duct (spgd); latter with swelling proximad spermathecal gland (spg). Without helminthoid sclerite. Without secondary spermathecal gland.

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).

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.

(Map 1). This species is known only from two montane localities in the Mexican state of Oaxaca.

Type specimens, only. For label details, see above.

Zuphium mexicanum Chaudoir, 1863 (here designated).

A compound Latinized noun, treated as neuter, from the generic name Zuphium and oides, resembling; hence meaning “resembling Zuphium”.

With character states of Western Hemisphere Zuphium genus-group, restricted as follows: size small, pronotum and dorsal surface of elytra densely setose, setae decumbent, body depressed, integument piceous to rufotestaceous, head capsule posteriorly relatively broad, laterally broadly rounded, antennae elongate, antennomere 1 as long or longer than antennomeres 2–4. Humeri broadly rounded. Metasternum long, metepisternum longer than wide at base. Macropterous. Male genitalia: phallus without dorsal paraostial sclerites (Fig. 10D–10F; cf. Fig. 13A–C, ps). Ovipositor: Gonocoxite 2 (Figs 11D–F; cf. Figs 11A–C, gc2) short, thick; in lateral aspect falciform, apex pointed; in dorso-ventral aspect, broad, paddle-like, apex broadly rounded, not notched. Female genital tract: without secondary spermathecal gland (Fig. 12B; cf. Fig. 13D, ssg).

None required here. See description of Zuphioides mexicanum, below.

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 Erwin (1991: 42), Erwin et al. (2012: 32), and Larochelle and Lariviere (2003: 514–515). Ball and Shpeley (2000: 397) mistakenly classified Zuphium as “xerophilous”, in part.

For Zuphioides mexicanum, two females, Oberthür-Chaudoir Collection, in front of the following box label: “mexicanum/ Chaud/ Mexique/ 57. Sallé”. LECTOTYPE female, labeled as follows: “33”/ “Ex Musaeo/Chaudoir” [red print]; “Lecto/type” [circular, ringed with red]; in Museum National d’Histoire Naturelle, Paris; PARALECTOTYPE labeled “ Ex Musaeo/Chaudoir” [red print]. Designated by Mateu (1985: 330). For Zuphium vicinum, holotype, in Institute of Zoology, Polish Academy of Sciences, Warsawa, #1711.

Indicated by Mateu (1985: 330) as Veracruz, for Zuphioides mexicanum; for Zuphium vicinum, “Mexico”.

A Latinized eponym, nominative case, based on the name of the country in which the type locality is located.

Size and proportions. Small, OBL 4.88–4.96 mm; EW 1.60–1.62 mm A1L/A2–4L 0.90–1.00; HW/PW 0.92–0.97. (For seven species of Zuphioides: OBL 4.64–6.586 mm; EW 1.60–1.84 mm A1L/A2–4L 0.88–1.12; HW/PW 0.91–0.97).

Color. Body rufotestaceous; appendages slightly paler; vestiture testaceous.

Habitus (Fig. 1B). Flat, overall. Head capsule (dorsal aspect, Fig. 2B) trapezoidal, obtusely angulate posteriolaterally.

Eyes (Figs 3B). Convex, easily seen, ommatidia clearly evident at 50× (about 16 ommatidia are crossed on a horizontal diameter).

Pronotum. Narrow.

Elytra (Fig. 1B). Lateral margins straight, not bowed, parallel to one another.

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.

Body vestiture and punctation. Dorsal surface of head and pronotum sparsely punctate, vestiture sparse. Elytra and abdominal sterna III-VII with punctation and vestiture dense, vestitural setae decumbent.

Fixed setae. Head (Fig. 4C): clypeus one pair; head capsule with anterior pair of supraorbitals (asos) above eyes; posterior pair of supraorbitals (psos) posteriad eyes; one pair of postocular setae (pos) immediately behind eyes, laterally; one pair of occipital setae (ocs) posteriorly and mediad eyes; posterior supernumerary setae lacking. Antennomere 1 (Figs 4B, 4D, as1) with single long seta distally, in addition to decumbent vestiture and row of small erect setae (as2, as3). Pronotum with two pairs of lateral marginal setae, anterior pair in anterior 1/8, posterior pair at posteriolateral angles. Prosternum with one pair setae anterioventrally (Fig. 9B). Elytra: each elytron anteriorly with parascutellar seta; lateral setae about 21 (one group anteriorly, one group posteriorly, and single seta medially. Abdominal sterna IV-VI without evident ambulatory setae, sternum VII with pair of long setae near posterior margin.

Head, dorsal aspect (Fig. 4C). Occiput posteriorly and postocciput markedly constricted, in form of narrow neck, postoccipital suture evident. Frontoclypeal suture present. Genal sulcus (gs) prominent, but less so than in Coarazuphium (cf. Fig. 4A).

Eyes (Fig. 4B). Macrophthalmous, markedly convex, readily seen, ommatidia evident at 50×, or lower magnification.

Antennae (Fig. 4D). Antennae filiform, extended about body length. Antennomere 1 rather slender, slightly longer than antennomeres a2-a4; antennomere 2 short, about half length of a3; antennomeres 3-11 narrow, cylindrical, distinctly longer than wide.

Mouthparts. As described for Zuphium genus-group, above. Labrum (Fig. 5C). Anterior margin concave. Epipharynx (Fig. 5D). Mandibles (Figs 7A–7F). Maxillae (Figs 8D, 8E). Labium, ventral aspect (Fig. 8F).

Pronotum (Fig. 1B). Anterior margin truncate, lateral margins markedly sinuate posteriorly, posteriolateral angles prominent, dentiform, slightly anteriad posterior margin; surface impressions (anterior and posterior transverse and median longitudinal) shallow; lateral grooves and posteriolateral impressions moderately deep.

Pterothorax. Metasternum of average length; metepisternum elongate, lateral margin longer than anterior margin.

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.

Hind wings. Macropterous; wings folded beneath elytra at rest.

Legs. Male fore-tarsus (Fig. 9A).

Abdomen. As described for Western Hemisphere Zuphium genus-group.

Male genitalia (Figs 10D–10F). Phallus narrow (PW/PL 0.254), slightly curved ventrally, narrowed apically, apical margin rounded, apical portion very short (AL/PL 0.036), ostial membrane extensive (OM/PL 0.317). Left paramere (lp) conchoid. Right paramere (rp) short, ovoid in form (RP/LP 0.317).

Female genitalia: ovipositor (Figs 11D–11F). Gonocoxite 1 (gc1) with patch of long trichoid setae distally on ventral surface. Gonocoxite 2 (gc2) short, thick; in lateral aspect falciform, apex pointed, row of four long trichoid setae dorso- and ventro-laterally; in dorso-ventral aspect, broad, paddle-like, apex broadly rounded, not notched; otherwise, as described for Western Hemisphere Zuphium genus-group.

Female genital tract (Fig. 12B). Bursa copulatrix (bc) bulbous (collapsed in Fig. 12B), inserted at base of common oviduct (co). Spermatheca (sp) inserted near junction of common oviduct and bursa copulatrix, markedly elongate and slender, with helminthoid sclerite (hs) at base. Spermathecal gland duct inserted near base of spermatheca, with distinct swelling proximad spermathecal gland (spg). Without secondary spermathecal gland.

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 (2012: 1357).