Hyboptera Chaudoir, 1872 of the Cryptobatida group of subtribe Agrina: A taxonomic revision with notes on their ways of life (Insecta, Coleoptera, Carabidae, Lebiini)

Abstract Hyboptera Chaudoir, 1872 of the Cryptobatida group of subtribe Agrina, Lebiini, living in the Neotropics and southernmost Nearctic realms are diagnosed, described, and illustrated, and new species are assigned to two inclusive species groups. Occurrences of species range from Texas, USA, to the state of Santa Caterina in Brazil. Seven new species of Hyboptera are described: Hyboptera biolat Erwin & Henry, sp. n.; Type locality – Perú, Madre de Dios, Manu Reserved Zone, Río Manu, BIOLAT Biological Station, Pakitza; Hyboptera vestiverdis Henry & Erwin, sp. n.; Type locality – Perú, Madre de Dios, Manu Reserved Zone, Río Manu, BIOLAT Biological Station, Pakitza; Hyboptera lucida Henry & Erwin, sp. n.; Type locality – French Guiana, Cayenne, Commune de Roura, Montagne des Chevaux; Hyboptera scheelea Erwin & Henry, sp. n.; Type locality – Perú, Loreto, Pacaya-Samiria National Reserve, Río Samiria (South Branch), Camp Terry; Hyboptera shasta Erwin, sp. n.; Type locality – Brazil, Amazonas, north of Manaus on Amazonas 010 at Km 26, Reserva Ducke; Hyboptera tepui Erwin & Henry, sp. n.; Type locality – Venezuela, Amazonas, Cerro de la Neblina, Río Baria Basecamp; Hyboptera tiputini Erwin & Henry, sp. n.; Type locality – Ecuador, Orellana, Yasuni National Park (edge), 95.43 km E (heading 101.46°) Coca, Tiputini Biodiversity Station. A revised identification key is provided to the genera of the Cryptobatida group and another to the species of Hyboptera Chaudoir and distributional data are provided for all known species of the latter. Adults of these species often occur in the canopy of many tropical tree species and records are reported where known. In addition, adults are found under the webbing of Psocoptera and in fleshy anther rings of Bombacaceae (Pseudobombax septenatum (Jacq.) Dugand), on the rain forest floor in the dry season. Larval and pupal stages of one species from Panamá are known from under bark of living fence posts; however, these immature stages are not treated in the current paper.


Introduction
The Cryptobatida Group of Lebiini was proposed to include a number of genera including Hyboptera Chaudoir (Erwin 2004). This genus was treated most recently by Reichardt (1971Reichardt ( , 1973 and those reports were brief and superficial, not full revisions. Before that, only five isolated species descriptions by Chaudoir, Reiche and Oberthür existed. Hence, this small group of very attractive and biologically interesting beetles requires an initial taxonomic revision. The purpose of this paper is to rapidly validate some new species names (Erwin and Johnson 2000) with descriptions and provide more complete re-descriptions of known species, so that the group is available for further study, particularly their way of life in association with Psocoptera. Hyboptera are members of the Cryptobatida group by virtue of the structure of their mouthparts, elytral transverse depression, and the male genitalia with a flagellum. Herein, we provide full descriptions of all known taxa, including color images, an up-to-date map of their known distributions, and what is known of their ways of life including host tree species.

Methods and materials
(modified from Erwin and Zamorano 2014) As noted in several past contributions, methods and species concepts follow those previously described (Erwin and Kavanaugh 1981;Kavanaugh and Erwin 1991). The species validation and diagnosis format follows as closely as possible that suggested in Erwin and Johnson (2000). For measurements, images of the specimens were taken using a Leica M420 stereoscope coupled to an EntoVision TM system. The resulting image was processed using the software Cartograph version 7.2.5 by Microvision Instruments. The magnification on the zoom was set to calibrate the system and it is embedded into the file of the image. The image was then opened with the software program Archimed version 6.1.4, also by Microvision, and the Measure tool was then used to determine the lengths of the various parts. A total of 239 images were obtained. Measurements of length (ABL, SBL) and width (TW) follow those of Ball (1972) and Kavanaugh (1979): ABL (apparent body length), measured from apex of labrum to apex of longer elytron (in adults of this genus, the abdomen often protrudes beyond the elytral apex, thus the ABL often is much larger that the SBL; SBL (standardized body length), equals the sum of the lengths of the head (measured from apex of clypeus to a point on midline at level of the posterior edge of compound eyes); PL (pronotum length) is measured from apical to basal margin along midline; LE (elytron length) is measured from apex of scutellum to apex of the longer elytron; TW (total width) is measured across both elytra at their widest point with suture closed; and WH (head width) is measured from extreme margin of protuberant eyes left to right. Note that not all specimens available were measured because more than 33 specimens were available, thus we limited "n" to 33 as a statistically valid sample size. Sexes were measured separately, we found slight differences among the species sexes, and hence we report measurements for both sexes in our Tables (see Appendix 1). For the Hyboptera treatment below, we provide relative size terms based on the SBL as follows: small-size < 3.3 mm, medium-size 3.3 mm to 4.3 mm, and large-size > 4.3 mm. For an explanation of the measurements and their incorporation in Appendix 1, see  and Erwin and Ball (2011). For the present study, we report the harmonic mean, as we believe it better reflects the central tendency than the arithmetic mean.
Attributes of the abdominal ventral sterna are referred to using the numbering system generally accepted in carabid studies, i.e., the sternum divided medially by the hind coxae is sternum II (the first being hidden) and the last visible is sternum VII (Liu et al. 2011). In a revision of the genus Pericompsus (Erwin 1974), a problem was encountered with the term "stria" for features of their punctate elytra (i.e., the so-called striae were not actually striae, rather they were rows of punctures). The result was the use of the term "interneur" to apply to the attribute lying between intervals. Through use of this term, one could describe the feature as interneur striate, punctate, striatopunctate, etc.
A similar problem exists for the proximal end of the median lobe of the male genitalia. In Snodgrass (1935), the term "phallobase" is used, and we have adopted it here (see . Therefore, by extension, in Carabidae, we can say phallobase hooded (e.g., Lebiini, Pseudomorphini), phallobase of two parallel sclerotized struts (basal trechines and Andinodontis), phallobase of two uneven struts (Bembidion), etc. Kavanaugh (pers. comm.) points out that with struts there are still connecting membranes surrounding the struts forming a "bulb." We have chosen the aedeagal illustration of a male H. angulicollis Chaudoir (Fig. 5A) and H. apollonia Erwin (Fig. 10A) to display the identifying code letters and these apply to all illustrations of male genitalia of Hyboptera included.
This study includes 738 adult specimens of Hyboptera, all currently at the National Museum of Natural History, Washington, DC (NMNH) and, where appropriate, returned to their corresponding institutions upon publication. Among these specimens, some were received from: Primary type specimens of new species will be deposited in their countries of origin if required by legal agreements, or museums of ownership at the conclusion of our studies on this tribe.

AMNH
The enhanced habitus images of the adult beetles portray most of the character states referred to in the key provided. Illustrations of male genitalia are standard for descriptive taxonomy of carabid beetles in both preparation and aspects presented, as is the presentation of the female genitalia (one example per genus, in this case H. lucida sp. n.). The habitus images of the adults were made with a Visionary Digital TM high resolution imaging system rendered using Photoshop to become "Digital Photoillustrations." Figure captions include an ADP number, which is a unique identification number for the specimen that was imaged and links the specimen and associated illustrations and/or images to additional information, such as collecting notes, in electronic databases at the NMNH.
Geographical data are presented for species based on all known specimens available at the time of manuscript preparation, including those in the literature. Georeferenced data have been determined from locality information provided on specimen labels; only those exact geo-references reported in decimal degrees that are provided on the label are placed in quotes. Otherwise, we have estimated others as closely as possible from places, mileage, or other locality data listed on the label and searched with Google Earth Pro. Latitude and longitude for those are reported in decimal degrees and have been corrected from those reported on the labels, if necessary; our bottom line is that georeferenced locality data reported herein are far more accurate than those provided on specimens labels.
A distribution map is provided for the species of Hyboptera (Fig. 11). Here, vernacular names in English are proposed, as common names are becoming increasingly needed in conservation reports and studies, and/or agricultural and forestry applications. These names are based on criteria set forth in Erwin (2011a) and applied in Erwin (2011b).
Host occurrences of rainforest trees are reported using the names provided by botanists who inventoried two fogging transects established by the senior author (TLE) in Ecuador. These names have not been elaborated with author names herein, as is traditional in botanical literature, however, they can be readily found on the internet.
Notes. Subtribe Agrina consists of those species formerly included in the Subtribe Calleidina (cf. Lorenz 2005). The Cryptobatida Group, by virtue of the attributes above, has the type genus Cryptobatis Eschscholtz. Many records of species in most genera refer to collections on fungus arrayed on sides or the bottom of fallen tree trunks. It is probable that the Cryptobatida Group contains many species whose adults are predators on other shelf fungus inhabitants (cf. Erwin and Erwin 1976, wherein adults and larvae of a species of Eurycoleus, which is a member of the subtribe Pericalina, are predaceous on fungal feeders; other pericalines are also associated with fungi). Hyboptera and its adelphotaxon Hybopteroides are exceptions (Erwin and Ball 2012), as the adults are likely predators on psocids and embiids beneath silken webs, respectively.
Below, we have added to the Cryptobatida Group the genus Straneotia Mateu 1961 by virtue of a study of new specimens collected by fogging and flight intercept traps (FIT) in Ecuador and French Guiana, respectively. Mateu erected this genus based on a single female specimen from eastern Amazonian Brazil (Straneotia freyi Mateu) and described a second species in the genus based on a single female from western Amazonian Brazil (Straneotia amazonica Mateu). These two species soon will be redescribed and the new species described (Erwin and Aldebron, in prep

Ways of life.
Much is known about the species in this genus and that is reported here for the first time. Adults of various species are regularly collected in both the wet and dry seasons using insecticidal fogging techniques in many species of trees reaching the forest canopy in the Amazon Basin, thus they are certainly mainly arboreal. They are good flyers as evidenced by their capture in Malaise traps, C.D.C. mosquito traps, FITs, and at different types of light traps. At Barro Colorado Island in Panamá, one of us (TLE) collected adults from the forest floor amongst the large shed anther rings of trees of the species Pseudobombax septenatum (Jacq.) Dugand; these rings being a moisture source on the forest floor during the dry season (Erwin 1991(Erwin , 2004. Warren E. Steiner collected several larvae and pupae along with emerging adults of Hyboptera verrucosa (Reiche) under bark of a living fence post in Panamá. These immature stages have yet to be described and illustrated (Erwin and Steiner, in prep).
Notes. Five species have been previously described in this genus, along with three species in its adelphotaxon, Hybopteroides Erwin & Ball, 2012. Apparently, Lorenz (1998 was unaware of the Reichardt (1973) paper designating a type species (H. angulicollis Chaudoir) for the genus and unnecessarily designated H. tuberculata (Dejean), as the type species. The former stands.
Included species. The species list below, as well as the arrangement of descriptions that follows, is ordered alphabetically within two species groups.

angulicollis species group
(recognized by Reichardt 1973) The most distinctive attribute of species in this group is that the pronotum has marked discal rugae etched almost horizontally and linear, as opposed to angulate. Adults of all have metallic green coloration somewhere on the dorsal surface and the general adult size is small to medium for the genus. Male phallus apex elongate, somewhat acuminate.
Derivation of specific epithet. The epithet, angulicollis, is a Latinized singular feminine adjective meaning "angled neck" referring to the angulate lateral sides of the pronotum.
Proposed english vernacular name. Angled-neck humps-backed beetle. Diagnosis. With the attributes of the genus and angulicollis species group as described above and adults with patches of bright metallic green para-medially on pronotum, elytra entirely with metallic green reflection, and venter entirely rufotestaceous.
Color: See diagnosis, above.
Prothorax: Pronotum moderately broad, disc centrally planar with dense transverse rugae. Lateral margins broadly explanate and obtusely angled medially then subtly arcuate to broadly obtuse hind angle, base medially produced and rounded posteriorly.
Pterothorax: Normal for Agrina, fully winged. Elytron intervals 3 with 6, and 5 with 5 (4) discal unisetiferous tubercles, other intervals moderately convex, side margin moderately explanate laterally only at middle third. Elytron broad and moderately short, moderately narrower than the pronotum at the broadest part, apex truncate, slightly rounded with distal corner broadly and obtusely rounded, disc not significantly convex, basal third slightly depressed. All interneurs well-impressed.
Legs: Femur dorso-ventrally moderately depressed, tibia coequal in length, more depressed; tarsus less than half the length of the tibia, fourth tarsomere markedly bilobed and with tarsal pad of setae.
Abdomen: Sparsely setiferous; normal ambulatory setae on sterna 3-5; female with two pairs of ambulatory setae on sternum 6, medial pair of setae less than the length of lateral pair; males with only the lateral pair of longer setae on sternum 6.
Female genitalia: Not investigated, likely similar to that of H. lucida (Fig. 8).
Variation: In the large series of specimens of this species available to us for this study from many localities, we noted that the pronotal explanate margins vary considerably in width, the more narrow they are the more prominent the midpoint angle, the broader the explanation the more obtuse the midpoint angle.
Dispersal potential. These beetles are macropterous and capable of flight. They are moderately swift and agile runners. Specimens have been acquired using most modern collecting methods, including insecticidal foggings, UV light sheets and traps, Malaise traps, SLAM Malaise traps, and flight intercept traps (FITs). Way of life. Adults are common in the lowlands (8 to 700 m a.s.l.) and appear to be generalists in a variety of rainforest biotopes including terra firme, várzea, igapó, second growth, subtropical moist and white sand forests. In these forests, they are commonly found in big trees with vines and epiphytes, in suspended dry leaves, under lianas close to the trunk, in dry Scheelea sp. and Astrocaryum chambira Burret palm fronds, in very large suspended dry Cecropia leaves, and in dry bamboo leaves of the genera Guadua [G. weberbaueri Pilg.] and Elytrostachys [sp.] Individuals can be found in all months of the year, in both the rainy and dry seasons. Member of this species have been recorded from the canopy of the following tree species using insecticidal fog-   Derivation of specific epithet. The specific epithet, biolat, is used as a noun in apposition based on the acronym of the Smithsonian Institution's past Program "Biodiversity in Latin America" (BIOLAT) which sought to field-train young Latin American biology students in biodiversity techniques and did so for over 200 of them between 1987 and 1991 in Perú and Bolivia. These beetles were collected under the auspices of the BIOLAT Program.
Proposed English vernacular name. Biolat humps-backed beetle. Diagnosis. With the attributes of the genus and angulicollis species group as described above and adults with patches of bright metallic green para-medially on pronotum, elytra entirely with metallic green patches, and venter with meso-and metathorax mostly infuscated.
Color: See diagnosis, above.
Prothorax: Pronotum moderately broad, disc centrally planar with dense transverse rugae. Lateral margins moderately explanate and obtusely angulate medially then straight to narrowly obtuse hind angle, base medially produced and rounded posteriorly.
Pterothorax: Normal for Agrina, fully winged. Elytron interval 3 with 7(8), and interval 5 with 5 (4) discal unisetiferous tubercles, other intervals moderately convex, side margin moderately explanate laterally only at middle third. Elytron broad and moderately short, moderately narrower than the pronotum at the broadest part, apex truncate, slightly rounded with distal corner broadly and obtusely rounded, disc not significantly convex, basal third slightly depressed. All interneurs well-impressed.
Legs: Femur dorso-ventrally moderately depressed, tibia coequal in length, more depressed; tarsus less than half the length of the tibia, fourth tarsomere markedly bilobed and with tarsal pad of setae.
Abdomen: Sparsely setiferous; normal ambulatory setae on sterna 3-5; female with two pairs of ambulatory setae on sternum 6, medial pair of setae less than the length of lateral pair; males with only the outer pair of longer setae on sternum 6.
Female genitalia: Not investigated, likely similar to that of H. lucida (Fig. 8).
Dispersal potential. These beetles are macropterous and probably capable of flight. They are moderately swift and agile runners. Specimens have been acquired using insecticidal fogging methods.
Way of life. Adults are common in the lowlands (356 m.a.s.l.) and appear to be generalists in a variety of rainforest biotopes including terra firme and upper floodplain forests. In these forests, they are commonly found in suspended dry leaves in Guadua weberbaueri Pilg. bamboo patches and among Astrocaryum chambira Burret palm dry leaf-skirts. Adults have been obtained in September-October; hence, they are active in the late dry and early rainy seasons.
Other specimens examined. Geographic distribution (Fig. 11). This species is currently known only from the type locality in Perú in the Río Manu watershed at the Pakitza Vigilante Post where the BIOLAT Biological Station operated from 1987 to 1992.
Notes. The holotype will be deposited in UNMSM and is currently held in trust until the completion of studies at NMNH. Specimen ADP007443 is missing its fore body. Derivation of specific epithet. The species epithet ''vestiverdis" is a Latinized feminine noun meaning green vest, referring to the two lobes of color on the pronotal disc on individuals of this species with central line of pale color bisecting the pattern and appearing as an open vest.

Hyboptera vestiverdis
Proposed English vernacular name. Leprechaun humps-backed beetle. Diagnosis. With the attributes of the genus and angulicollis species group as described above and adults with only dark slightly-metallic markings on the pronotal disc, elytra olivaceous, markedly shiny, some individuals with hint of metallic green near basal margin, narrow, with slightly arcuate lateral margin.
Color: See diagnosis, above.
Pterothorax: Normal for Agrina, fully winged. Elytron intervals 3 with 7 and 5 with 5(6) elongate unisetiferous tubercles, other intervals moderately convex, side margin broadly explanate laterally only at middle third. Elytron broad and short, slightly narrower than the pronotum at the broadest part, apex truncate, slightly rounded with distal corner broadly and obtusely rounded, disc not significantly convex, basal third slightly depressed. All interneurs well-impressed.
Legs: Femur dorso-ventrally moderately depressed, tibia coequal in length, more depressed; tarsus less than half the length of the tibia, fourth tarsomere markedly bilobed and with tarsal pad of setae.
Abdomen: Sparsely setiferous; normal ambulatory setae on sterna 3-5; female unknown; males with only the lateral pair of longer setae on sternum 6.
Female genitalia: Not investigated, likely similar to that of H. lucida (Fig. 8). Dispersal potential. These beetles are macropterous and capable of flight. They are moderately swift and agile runners. Specimens have been acquired in Malaise traps and using insecticidal fogging methods.
Way of life. Adults are common in the lowlands (116 to 356 m.a.s.l.) and appear to be generalists in a variety of rainforest biotopes including terra firme and várzea. In these forests, they are commonly found in big trees with vines and epiphytes, in suspended dry leaves, in dry Scheelea sp. and Astrocaryum chambira Burret palm frond skirts, in dry leaves scattered in vine tangles, and in dry leaves of the bamboo Guadua weberbaueri Pilg. Individuals can be found in January-February, May-July, and September-October, in both the rainy and dry seasons. Member of this species have been recorded from the canopy of the following tree species using insecticidal fogging techniques: Grias neuberthii; Eschweilera  Geographic distribution (Fig. 11). This species is currently known from the western Amazon Basin in lowland Ecuador and Perú. Notes. The holotype will be deposited in UNMSM and is currently held in trust until the completion of studies at NMNH. Derivation of specific epithet. The species epithet "scheelea" is used as a noun in apposition which is based on the genus of palm upon which the holotype was found.

Hyboptera scheelea
Proposed English vernacular name. Palm-frond humps-backed beetle. Diagnosis. With the attributes of the genus and angulicollis species group as described above and adults without pronotal markings, pronotal surface rufotestaceous, elytron blackish-blue with metallic green highlights across the humeri and green points at some larger tubercles with lateral margin narrowly testaceous to latero-apical corner but not reaching sutural apex. Apical abdominal tergite mostly infuscated with narrow median testaceous stripe. Size smaller than H. shasta adults.
Prothorax: Pronotum markedly broad, disc centrally depressed with dense transverse rugae. Lateral margins broadly explanate and obtusely angulate medially then moderately arcuate to obtuse hind angle, base medially produced and rounded posteriorly.
Pterothorax: Normal for Agrina, fully winged. Elytron intervals 3 and 5 each with (4)5 discal unisetiferous tubercles, interval 3 with one such tubercle near apex, other intervals moderately convex, side margin broadly explanate laterally only at middle third. Elytron broad and short, slightly narrower than the pronotum at the broadest part, apex truncate, slightly rounded with distal corner broadly and obtusely rounded, disc not significantly convex, basal third slightly depressed. All interneurs well-impressed.
Legs: Femur dorso-ventrally moderately depressed, tibia coequal in length, more depressed; tarsus less than half the length of the tibia, fourth tarsomere markedly bilobed and with tarsal pad of setae.
Abdomen: Sparsely setiferous; normal ambulatory setae on sterna 3-5; female with two pairs of ambulatory setae on sternum 6, medial pair of setae less than the length of lateral pair; males unknown.
Male genitalia: Unknown. Female genitalia: Not investigated, likely similar to that of H. lucida (Fig. 8). Dispersal potential. These beetles are macropterous and probably capable of flight. They are moderately swift and agile runners. The holotype was acquired by insecticidal fogging of a Sheelea palm.
Way of life. The single known adult was found in May in lowlands (129 m.a.s.l.) in the secondary floodplain of igapó forests.
Other specimens examined. None. Geographic distribution (Fig. 11). This species is currently known only from the type locality in the lowland of Amazonian Perú.
Notes. The holotype will be deposited in UNMSM and is currently held in trust until the completion of studies at NMNH. Derivation of specific epithet. The species epithet "shasta" is an eponym based on the first name of the coauthor of this paper and former Intern in the laboratory of the senior author of this paper at the Smithsonian Institution. Proposed English vernacular name. Shasta's humps-backed beetle. Diagnosis. With the attributes of the genus and angulicollis species group as described above and adults without pronotal markings, pronotal surface rufotestaceous, elytron brilliant metallic green throughout with lateral margin testaceous to latero-apical corner, not quite reaching sutural apex; apical abdominal tergite mostly infuscated with narrow median testaceous stripe; size larger than H. scheelea adults.
Pterothorax: Normal for Agrina, fully winged. Elytron intervals 3 with 7 and 5 with 3/4 elongate unisetiferous tubercles, other intervals moderately convex, side margin broadly explanate laterally only at middle third. Elytron broad and short, slightly narrower than the pronotum at the broadest part, apex truncate, slightly rounded with distal corner broadly and obtusely rounded, disc not significantly convex, basal third slightly depressed. All interneurs well-impressed.
Legs: Femur dorso-ventrally moderately depressed, tibia coequal in length, more depressed; tarsus less than half the length of the tibia, fourth tarsomere markedly bilobed and with tarsal pad of setae.
Abdomen: Sparsely setiferous; normal ambulatory setae on sterna 3-5; female unknown; males with only the lateral pair of longer setae on sternum 6.
Male genitalia: Not investigated as only the holotype male is available, likely similar to that of H. angulicollis (Fig. 5A).
Female genitalia: Not investigated, likely similar to that of H. lucida (Fig. 8).
Dispersal potential. These beetles are macropterous and capable of flight. They are moderately swift and agile runners. The holotype was acquired in a C.D.C. light trap.
Way of life. The single known adult was found in July in lowlands (70 m.a.s.l.) in the terra firme forests.
Other specimens examined. None. Geographic distribution (Fig. 11). This species is currently known only from the type locality in lowland Amazonian Brazil. Derivation of specific epithet. The species epithet "tepui" is used as a noun in apposition and is based on the type of Venezuelan flat-topped upland near which the holotype was found.

Hyboptera tepui
Proposed English vernacular name. Tepui humps-backed beetle. Diagnosis. With the attributes of the genus and angulicollis species group as described above and adults without pronotal markings, pronotal surface rufotestaceous. Elytron with lateral margin broadly testaceous from humerus to sutural apex; apical abdominal tergite testaceous with slight infuscation at extreme posterior-lateral corners.
Pterothorax: Normal for Agrina, fully winged. Elytron intervals 3 and 5 each with 6 discal unisetiferous tubercles, interval 3 with one subtle tubercle near apex, other intervals shallowly to moderately convex, side margin broadly explanate laterally only at middle third. Elytron broad and short, slightly narrower than the pronotum at the broadest part, apex truncate, slightly rounded with distal corner broadly and obtusely rounded, disc not significantly convex, basal third slightly depressed. All interneurs well-impressed.
Legs: Femur dorso-ventrally moderately depressed, tibia coequal in length, more depressed; tarsus less than half the length of the tibia, fourth tarsomere markedly bilobed and with tarsal pad of setae.Abdomen: Sparsely setiferous; normal ambulatory setae on sterna 3-5; female with two pairs of ambulatory setae on sternum 6, medial pair of setae less than the length of lateral pair; males unknown.
Male genitalia: Unknown. Female genitalia: Not investigated, likely similar to that of H. lucida (Fig. 8). Dispersal potential. These beetles are macropterous and capable of flight. They are moderately swift and agile runners. The holotype was acquired at black light in a rainforest clearing.
Way of life. The single known adult was found in February in lowland (138 m.a.s.l.) terra firme forests.
Other specimens examined. None. Geographic distribution (Fig. 11). This species is currently known only from the type locality in lowland Venezuela.  Derivation of specific epithet. The specific epithet, "tiputini" is used as a noun in apposition and is based on the Tiputini Biodiversity Station in the Yasuni area of eastern Ecuador in reference to one of the places in which members of this species are found.

Hyboptera tiputini
Proposed English vernacular name. Tiputini humps-backed beetle. Diagnosis. With the attributes of the genus and angulicollis species group as described above and adults with patches of bright metallic green para-medially on pronotum, elytra markedly shiny, violaceous, some individuals with hint of metallic green near basal margin, broad, with markedly arcuate lateral margin.
Color: See diagnosis, above.
Pterothorax: Normal for Agrina, fully winged. Elytron interval 3 with 7(8) and interval 5 with 5 unisetiferous tubercles, other intervals shallowly to moderately convex, side margin broadly explanate laterally only at middle third. Elytron broad and short, slightly narrower than the pronotum at the broadest part, apex truncate, slightly rounded with distal corner broadly and obtusely rounded, disc not significantly convex, basal third slightly depressed. All interneurs well-impressed.
Legs: Femur dorso-ventrally moderately depressed, tibia coequal in length, more depressed; tarsus less than half the length of the tibia, fourth tarsomere markedly bilobed and with tarsal pad of setae.
Abdomen: Sparsely setiferous; normal ambulatory setae on sterna 3-5; female with two pairs of ambulatory setae on sternum 6, medial pair of setae less than the length of lateral pair; males unknown.
Female genitalia: Not investigated, likely similar to that of H. lucida (Fig. 8). Dispersal potential. These beetles are macropterous and probably capable of flight. They are moderately swift and agile runners.
Way of life. Adults are common in the lowlands and lower midlands (20 to 900 m a.s.l.) and appear to be generalists in a variety of rainforest biotopes including terra firme, várzea, and igapó. In these forests, they are commonly found in big trees with vines and epiphytes, in suspended dry leaves, in dry Astrocaryum chambira Burret palm frond skirts. Individuals can be found in January-November, in both the rainy and dry seasons. Member of this species have been recorded from the canopy of the following tree species using insecticidal fogging techniques: Geographic distribution (Fig. 11). This species is currently known from the type locality and other localities in Ecuador and various localities in Colombia and Perú.
Notes. The holotype will be deposited in Escuela Politécnica Nacional, Quito, Ecuador and is currently held in trust until the completion of studies at NMNH.
Derivation of specific epithet. The specific epithet, viridivittis, is a feminine Latin adjective referring to the green line pattern of the pronotum.
Proposed English vernacular name. Green-lined humps-backed beetle. Diagnosis. With the attributes of the genus and angulicollis species group as described above and adults with patches of bright metallic green para-medially on pronotum, elytra dark matte black, some individuals with hint of metallic green near basal margin. Venter with gular region, prosternal region, meso-and metathroax, and margins of abdominal sterna infuscated, otherwise rufous.
Description. (Figs 4B, 5E). Size: See Appendix 1. Length (SBL) medium for ge-Notes. Reichardt (1971Reichardt ( , 1973 reported the following additional specimens that we did not see: Brazil -Minas Gerais: no locality (2 exs. MNHP). Rio de Janeiro: Nova Friburgo (l ex. BMNH). Guanabara, (5 exs. BMNH, MNHP, MZSP). São Paulo: Barueri (l ex. MZSP). Santa Catarina: Nova Teutonia (1 ex. BMNH), Corupá (1 ex. MCZ). Additionally, Chaudoir (1872) mentioned he had two specimens; therefore, a lectotype needs to be designated. Reichardt (1973) mistakenly writes he saw the "holotype." We have chosen the first of Chaudoir's specimens as the Lectotype (see above). tuberculata species group (recognized by Reichardt 1973) The most distinctive attribute of species in this group is that the pronotum has discal rugae etched at an angle aimed medio-posteriorly, or somewhat chaotically seemingly without order. Adults of all species have no metallic coloration (except H. apollonia with subtle traces only) on the dorsal surface and the general adult size is medium to large for the genus. Male phallus apex short, broadly or narrowly blunt.
Derivation of specific epithet. The specific epithet, apollonia, is an eponym based on the first name of Michael Corleone's beautiful young Italian wife in the movie The Godfather whose death in a car explosion perpetrated by Mafia competition signifies the useless instantaneous death of so many species when humans put fire to the tropical rain forest in time of drought.
Proposed English vernacular name. Apollonia's humps-backed beetle. Diagnosis. With the attributes of the genus and angulicollis species group as described above and with only dark non-metallic markings on the pronotal disc; pronotum with discal rugae etched horizontally and linear. Elytra black with paler margin not reaching suture. Mouthparts, appendages, margin of prothorax, venter of head and prothorax, abdominal segments II-V testaceous; meso-and metathorax, and abdominal segment VI infuscated.
Prothorax: Pronotum markedly broad, disc centrally slightly depressed with dense transverse rugae. Lateral margins very broadly explanate and obtusely rounded medially then nearly straight to obtuse hind angle, base medially produced and rounded posteriorly.
Pterothorax: Normal for Agrina, fully winged. Elytron intervals 3 with 7, and interval 5 with 6 (5) discal unisetiferous tubercles, other intervals moderately convex, side margin moderately explanate laterally only at middle third. Elytron broad and moderately short, moderately narrower than the pronotum at the broadest part, apex truncate, slightly rounded with distal corner broadly and obtusely rounded, disc not significantly convex, basal third slightly depressed. All interneurs well-impressed.
Legs: Femur dorso-ventrally moderately depressed, tibia coequal in length, more depressed; tarsus less than half the length of the tibia, fourth tarsomere markedly bilobed and with tarsal pad of setae.
Abdomen: Sparsely setiferous; normal ambulatory setae on sterna 3-5; female with two pairs of ambulatory setae on sternum 6, medial pair of setae less than the length of lateral pair; males with only the lateral pair of longer setae on sternum 6.
Female genitalia: Not investigated, likely similar to that of H. lucida (Fig. 8). Dispersal potential. These beetles are macropterous and capable of flight. They are moderately swift and agile runners. Modern collecting methods have acquired specimens, including insecticidal foggings and malaise traps.
Way of life. An adult of this species was fogged from a tree in the genus Guarea at La Selva and another caught in a Malaise trap as part of the ALAS Project. F. Nevermann collected a specimen on a rotten log. The known elevational range of this species is between 9 and 815 m.a.s.l. Adults have been obtained in January-April, and September-October; hence they are active in both the dry and rainy seasons in the lowlands and lower middle altitudes, both on the east and west sides of the Cordillera Central.
Other Geographic distribution (Fig. 11). This species is currently known from the type locality in Panamá and throughout southern Central America north to Costa Rica.
Notes. Adults of this species are unusual in that they subtly bear two attributes similar to species in the angulicollis species-group: faint traces of metallic green and a short narrow nubnen-like apex of the phallus, rather than short and broadly blunt. Adults of the angulicollis species-group are extensively green and the phallus apex, while narrow is elongate.
Derivation of specific epithet. The specific epithet, auxiliadora, is an eponym based on the middle name of María Auxiliadora Sanchez, who for many years was responsible for the welfare of participating visiting taxonomists at INBio and its facilities and who made life easy therein while we undertook our studies of the rich Costa Rican fauna and flora.

Diagnosis.
With the attributes of the genus and tuberculata species group as described above and adults with only dark non-metallic markings on the pronotal disc, venter substantially infuscated, pronotum with discal rugae etched at an angle aimed medio-posteriorly, or somewhat chaotically; elytron with sutural margin at apical sixth pale, not black, if brownish not contrasting with background color, and elytra just posterior to scutellum with a V-shaped pale area encompassing the sutures and first intervals. Elytron broad with markedly arcuate lateral margin.
Prothorax: Pronotum markedly broad, disc centrally depressed along midline with dense arcuate rugae directed anteriorly in apical half, posteriorly in posterior half of disc. Lateral margins broadly explanate and subtly produced at lateral seta, but not acute medially then straight to obtuse hind angle, base medially slightly produced and rounded posteriorly.
Pterothorax: Normal for Agrina, fully winged. Elytron interval 3 with 10 discal unisetiferous tubercles and interval 5 with 9(8) discal unisetiferous tubercles, interval 3 with one such tubercle near apex, other intervals moderately convex, side margin broadly explanate laterally at middle third. Elytron moderately broad and short, moderately narrower than the pronotum at the broadest part, apex truncate, slightly rounded with distal corner broadly and obtusely rounded, disc not significantly convex, basal third slightly depressed. All interneurs well-impressed.
Legs: Femur dorso-ventrally moderately depressed, tibia coequal in length, more depressed; tarsus less than half the length of the tibia, fourth tarsomere markedly bilobed and with tarsal pad of setae.
Abdomen: Sparsely setiferous; normal ambulatory setae on sterna 3-5; female with two pairs of ambulatory setae on sternum 6, medial pair of setae less than the length of lateral pair; males with only the lateral pair of longer setae.
Female genitalia: Not investigated, likely similar to that of H. lucida (Fig. 8).  Veracruz, 3.58 km NE Catemaco, Lake Catemaco, 18.444 N, 95.078 W, 355m, 23 August 1967 (GE Ball, TL Erwin, et al.) Geographic distribution (Fig. 11). This species is currently known only from the type locality and nearby areas in Texas, and from Panamá, and in between those extremes only from México, Honduras, and Costa Rica. It likely is to be found in other Central American countries with further sampling.
Derivation of specific epithet. The specific epithet, dilutior, is an adjective referring to the "washed out" appearance of the color of adults of this species in comparison with those of other species.
Proposed English vernacular name. Oberthür's humps-backed beetle. Diagnosis. With the attributes of the genus and tuberculata species group as described above and adults with only dark non-metallic markings on the head and pronotal disc, pronotum with discal rugae in basal half etched at an angle aimed medioposteriorly, or somewhat chaotically. Elytra mostly testaceous with darkly marked tubercles; small medio-apical tubercles also infuscated; with sutural margin at apical sixth infuscated markedly contrasting with testaceous background color. Venter of head and thorax substantially infuscated; abdominal sterna mostly pale with subtle infuscation.
Color: See diagnosis, above.
Prothorax: Pronotum markedly broad, disc centrally markedly depressed with coarse angulate rugae. Lateral margins broadly explanate and evenly rounded to obtuse hind angle, base medially produced and rounded posteriorly.
Pterothorax: Normal for Agrina, fully winged. Elytron intervals 3 and 5 each with 7 promient discal unisetiferous tubercles, other intervals moderately convex, side margin broadly explanate laterally only at middle third. Elytron broad and short, much narrower in width to that of the pronotum at the broadest part, apex truncate, slightly rounded with distal corner broadly and obtusely rounded, disc not significantly convex, basal third slightly depressed. All interneurs well-impressed.
Legs: Femur dorso-ventrally moderately depressed, tibia coequal in length, more depressed; tarsus less than half the length of the tibia, fourth tarsomere markedly bilobed and with tarsal pad of setae.
Abdomen: Sparsely setiferous; normal ambulatory setae on sterna 3-5; female with two pairs of ambulatory setae on sternum 6, medial pair of setae less than the length of lateral pair; males with only the lateral pair of longer setae. Male genitalia: Phallus ( Fig. 10B) with ostium of 1/6 its length, catopic, apex very short, narrowly pointed, broadly rounded in dorsal aspect; endophallus with flagellum (obvious in illustration), flagellum not barbed. Parameres asymmetric, right very small, left larger.
Female genitalia: Not investigated, likely similar to that of H. lucida (Fig. 8). Dispersal potential. These beetles are macropterous and capable of flight. They are moderately swift and agile runners. Adults of this species are attracted to C.D.C. light traps and captured in SLAM Malaise traps.
Way of life. Adults are common in the lowlands (7 to 356 m a.s.l.) and appear to be generalists in a variety of rainforest biotopes including terra firme and várzea. In these forests, they are commonly found in big trees with vines and epiphytes, in suspended dry leaves, in dry Sheelea sp. and Astrocaryum chambira Burret palm frond skirts. Individuals can be found from January to December, in both the rainy and dry seasons. Member of this species have been recorded from the canopy of the following tree species using insecticidal fogging techniques: Chrysophyllum argenteum cf.; Sterculia colombiana cf.; Parkia multijuga cf.; Naucleopsis herrerensis cf.; Matisia malacoca- Proposed english vernacular name. Pied humps-backed beetle. Diagnosis. With the attributes of the genus and tuberculata species group as described above and adults with only dark non-metallic markings on the pronotal disc; elytra black with 4 small pale medio-apical tubercles. Venter completely piceous. Largest adults in the genus. Description. (Figs 7B,8,10C). Size: See Appendix 1. Length (SBL) long for genus, ABL = 5.51-6.67 mm, SBL = 4.38-5.27 mm, TW = 2.36-3.25 mm.
Color: See diagnosis, above.
Prothorax: Pronotum markedly broad, disc centrally shallowly depressed with coarse angulate rugae. Lateral margins broadly explanate and evenly rounded to level of lateral seta then straight to obtuse hind angle, base medially produced and rounded posteriorly.
Pterothorax: Normal for Agrina, fully winged. Elytron intervals 3 with 10 and interval 5 with 7 prominent discal unisetiferous tubercles, interval 4 with sub-apical pale colored "lens," other intervals moderately convex, side margin broadly explanate laterally only at middle third. Elytron broad and moderately short, much narrower in width than that of the pronotum at the broadest part, apex truncate, slightly rounded with distal corner broadly and obtusely rounded, disc not significantly convex, basal third slightly depressed. All interneurs well-impressed.
Legs: Femur dorso-ventrally moderately depressed, tibia coequal in length, more depressed; tarsus less than half the length of the tibia, fourth tarsomere markedly bilobed and with tarsal pad of setae.
Abdomen: Sparsely setiferous; normal ambulatory setae on sterna 3-5; female with two pairs of ambulatory setae on sternum 6, medial pair of setae less than the length of lateral pair; males with only the lateral pair of longer setae.
Dispersal potential. These beetles are macropterous and capable of flight. They are moderately swift and agile runners. Adults of this species are attracted to light traps, and have been collected in FIT, Malaise, and SLAM traps.
Way of life. Adults are found in January-March, May-December, in all seasons, in lowlands (13-325 m.a.s.l.) in the lowland forests of Guyane.
Other specimens examined. Geographic distribution (Fig. 11). This species is currently known from the type locality in French Guiana and nearby areas.
Derivation of specific epithet. The species epithet ''tuberculata'' is a Latinized singular feminine adjective, referring to the bumpy attributes of the elytra.
Proposed English vernacular name. Tuberculate humps-backed beetle. Diagnosis. With the attributes of the genus and tuberculata species group as described above and adults with only dark non-metallic markings on the pronotal disc, elytron just posterior to scutellum with only the suture pale in color, otherwise markedly infuscate.
Pterothorax: Normal for Agrina, fully winged. Elytron intervals 3 with 8(9) and interval 5 with 6(7) prominent discal unisetiferous tubercles, interval 4 and 6 with subapical pale colored "lens," other intervals moderately convex, side margin broadly explanate laterally only at middle third. Elytron broad and moderately short, moderately narrower in width than that of the pronotum at the broadest part, apex truncate, slightly rounded with distal corner broadly and obtusely rounded, disc not significantly convex, basal third slightly depressed. All interneurs well-impressed.
Legs: Femur dorso-ventrally moderately depressed, tibia coequal in length, more depressed; tarsus less than half the length of the tibia, fourth tarsomere markedly bilobed and with tarsal pad of setae.
Abdomen: Sparsely setiferous; normal ambulatory setae on sterna 3-5; female with two pairs of ambulatory setae on sternum 6, medial pair of setae less than the length of lateral pair; males with only the outer pair of longer setae. Male genitalia: Phallus (Fig. 10D) robust with ostium of 1/6 its length, catopic, apex very short, broadly rounded in dorsal and lateral aspects; endophallus with flagellum (obvious in illustration), flagellum not barbed. Parameres asymmetric, right very small, left larger.
Female genitalia: Not investigated, likely similar to that of H. lucida (Fig. 8). Dispersal potential. These beetles are macropterous and capable of flight. They are moderately swift and agile runners. Adults of this species are attracted to C.D.C. Light traps.
Way of life. Adults are common in the lowlands and lower midlands (7-914 m a.s.l.) and appear to be generalists in a variety of rainforest biotopes including terra firme, dry and humid tropical forests, in gallery forests, and savanna forest patches. In these forests, they are commonly found in big trees with vines and epiphytes, in suspended medium to large-sized dry leaves, and in dense vine tangle plus leaves with light bamboo occurrence. Individuals can be found from January to December, in both the rainy and dry seasons. Member of this species have been recorded from the canopy of the following tree species using insecticidal fogging techniques:  Reichardt (1971: 53).
Derivation of specific epithet. The specific epithet, verrucosa, is a feminine Latin adjective referring to the tuberculate elytra.
Proposed English vernacular name. Verrucose humps-backed beetle. Diagnosis. With the attributes of the genus and tuberculata species group as described above and adults with only dark non-metallic markings on the pronotal disc, pronoum with rounded margins, elytron just posterior to scutellum with a V-shaped pale area encompassing the scutellum and sutural and first interval, venter substantially infuscate, and elytra narrow, not broadly arcuate.
Color: See diagnosis, above. Luster: Without any metallic highlights, shiny and matte. Microsculpture: Mostly isodiametric and stretched, shallowly impressed, cells somewhat more stretched around elytral tubercles.
Prothorax: Pronotum moderately broad, disc centrally depressed with coarse angulate rugae. Lateral margins broadly explanate and evenly rounded to obtuse hind angle, base medially produced and rounded posteriorly.
Pterothorax: Normal for Agrina, fully winged. Elytron intervals 3 with 8, and interval 5 with 6 prominent discal unisetiferous tubercles, interval 4 with sub-apical pale colored "lens," other intervals moderately convex, side margin broadly explanate laterally only at middle third. Elytron broad and moderately short, much narrower in width than that of the pronotum at the broadest part, apex truncate, slightly rounded with distal corner broadly and obtusely rounded, disc not significantly convex, basal third slightly depressed. All interneurs well-impressed.
Legs: Femur dorso-ventrally moderately depressed, tibia coequal in length, more depressed; tarsus less than half the length of the tibia, fourth tarsomere markedly bilobed and with tarsal pad of setae. mouthparts mentioned above, with Hyboptera adults, such as a serial row of long setae on three or more elytral intervals, angulate (or, subangulate) lateral margin of the pronotum, short antennae, broadly depressed mandibles, etc.
With regard to the genus Hyboptera, the recent discoveries of several new species in remote parts of the upper Amazon Basin suggests that further sampling in such areas will increase the species richness of this markedly (structurally and behaviorally) interesting (architecturally and behaviorally) lineage of Carabidae. We also note that even though at present there are not many species known and adults are morphologically markedly modified from more "typical" carabids, and have a unique way of life preying on insects under webbing, the lineage is widely dispersed from Texas to southeastern Brazil with many species that are widespread in their distributions.
Adults of the (currently) monobasic Thoasia Liebke, 1939 are exceedingly common in canopy fogging samples (Erwin 1991); however, nothing is known about their way of life and they are only known with precise location from foggings in Perú and Ecuador and FIT samples in French Guiana. Feeding specializations such as those hypothesized herein for adult Hyboptera and Hybopteroides and commonality of morphological attributes offer a fertile field of study on Thoasia for coleopterists eager to spend long periods of time in the rainforest canopies. However, before that, Thoasia is in need of a taxonomic revision and three undescribed species need to be treated (cf. Erwin, in prep). Liebke's holotype of the type species, Thoasia rugifrons, is in the Polish Academy of Sciences Collection according to Mroczkowski (1960).

Appendix 1
Morphological measurements and ratios for adults of species of Hyboptera Chaudoir 1872. All values are in millimeters. Apparent body length (ABL) is also provided in the descriptions. Means provided for ratios are "harmonic means."