taxonID	type	description	language	source
795187DDFFCDEF05F4D5D76FEE03E18D.taxon	description	As outlined in Nabozhenko & Steiner (2021: 354 - 356), the new world “ Helops ” clearly do not belong to the Old-World genus Helops as both belong to different putative phylogenetic groups. Mediterranean Helops to the Helopioid Helopina and New World “ Helops ” to the Catomoid Helopina. Horn (1870) recognized the single genus Helops for all Helopini from north of Mexico, which has remained constant until the last decade. Champion (1887) treated the Central American fauna and recognized three genera (See Nautes and Tarpela below) and retained the use of Helops for flightless species. Champion’s concept of a separate flightless genus would likely have included Stenotrichus LeConte, 1862 and Biomorphus Motschulsky, 1872 from the western United States which are both currently synonymized under Helops following the broad concept of Nearctic workers. Regardless of whether a broader or narrower concept is used, and whether flight is a meaningful character to separate genera, the genus name Helops is insufficient to represent the New World species currently placed there. The necessity of separating the Palearctic true Helops from the New World “ Helops ” has been discussed many times by previous authors: (Gebien 1943, Marcuzzi 2001, Dajoz 2001, Nabozhenko et al. 2016, Nabozhenko & Steiner 2021). Dinomus, as discussed above, agrees with Champion’s concept of flightless Central American species. This represents the oldest available genus-group name of Helopini with a type species from in the New World from which all other New World genera must be adequately distinguished from if they are to continue to be recognized.	en	Aalbu, Rolf L., Johnston, M. Andrew (2025): Reorganization of New World Helopini (Coleoptera: Tenebrionidae: Tenebrioninae). Zootaxa 5631 (1): 103-120, DOI: 10.11646/zootaxa.5631.1.4, URL: https://doi.org/10.11646/zootaxa.5631.1.4
795187DDFFCCEF05F4D5D22BE88EE36D.taxon	description	Dajoz (2001) recognized that New World “ Helops ” were different from Mediterranean Helops. He proposed the genus Neohelops describing the species Neohelops texanus Dajoz (Fig. 2) from the Big Bend region in Texas, USA. Though his species appears valid, his generic concept gives no putative diagnostic characters which separate this species from any other forms from the United States or Central America.	en	Aalbu, Rolf L., Johnston, M. Andrew (2025): Reorganization of New World Helopini (Coleoptera: Tenebrionidae: Tenebrioninae). Zootaxa 5631 (1): 103-120, DOI: 10.11646/zootaxa.5631.1.4, URL: https://doi.org/10.11646/zootaxa.5631.1.4
795187DDFFCCEF00F4D5D6FBE9E9E1A9.taxon	description	Nabozhenko & Steiner (2021) described a new genus based on three flightless species from the southeastern United States: Doyenellus cisteloides (Germar); and two new species, Doyenellus worthleyorum (Steiner) and Doyenellus theodorei (Steiner & Nabozhenko). Doyenellus is the only recently and comprehensively circumscribed genus of New World Helopina which allows for a more detailed examination of putatively diagnostic character states than the rest of the genera discussed herein. The following six characters were reported as unique or diagnostic of Doyenellus which are worth further examination. 1. The “ eighth elytral interval convex or keel-shaped and connected with elytral margin apically ” (Steiner & Nabozhenko 2021: 571). The eighth elytral interval (with the interval between the elytral fusion and the first stria counting as interval one and counting outward towards the epipleuron) is variably developed among the three species presently placed in Doyenellus and is particularly weakly elevated in D. theodorei (Fig. 3 A). There is large variation in the general convexity, flatness, and punctation of elytral intervals across Helopina species in North America. A keeled or convex eighth elytral interval is found in a number of species, including Helops rufipes (LeConte) (Fig. 3 F). 2. Male venter with “ elevated male abdominal ventrites 3 – 4, enlarged intersegmental membranes and strongly convex abdominal ventrite 5 (unique sexual dimorphism within the subtribe Helopina) ” (Nabozhenko & Steiner 2021: 571). The secondary sexual characters of the male abdomen are quite variable among New World Helopina, especially with regard to the fifth ventrite which is often inflated or depressed and can bear a wide range of setose fields or patches. The reportedly unique structure of the enlarged intersegmental membranes and convex fifth ventrite is also found in other species, including Tarpela micans (Fabricius, 1798) (Fig. 3 B), likely the most commonly collected species in the eastern United States. Moreover, this character is highly variably developed in the three species originally included in Doyenellus (see figures 13, 15, 17 in Nabozhenko & Steiner 2021: 569). This character is not only not unique among the New World fauna, but the variation seen in other species currently assigned among the genera Tarpela and Helops lead us to believe these characters are best used for species-level diagnoses. 3. Males further characterized by “ reduced, delimited and weakly sclerotized auriform apices of male inner sternite VIII (unique character within Helopini) ” (Nabozhenko & Steiner 2021: 571). As with the external secondary sexual characters discussed above, male sternite VIII displays a large amount of variation across New World Helopina species. In all species examined by us, this sternite is deeply emarginate medially, separating the distal half of the sclerite into a pair of sublateral lobes. These lobes are typically less sclerotized and appear more fleshy than the proximal portion of the sclerite. The lobes range from large and paddle shaped, to narrowly pointed triangles (Fig. 3 E), and to elongated almost digitate processes. The state defined for Doyenellus is not unique or easily characterized as a discrete state among the rest of the variation in this fauna. Moreover, the “ auriform ” lobes given as a generic character for Doyenellus are most distinct in D. cisteloides and are quite different in the other two species (see figures 34, 42, 51 in Nabozhenko & Steiner 2021). We similarly find this character useful for diagnosing particular species but the variation in the species presently placed in Doyenellus merely reflects some of the variation seen across New World Helopina and is not satisfactory for delimiting a genus. 4. Male penis (or median lobe) “ having a dorsally pubescent, narrowly rounded apex and ventral bifurcated bare apex ” as opposed to the “ typical catomoid median lobe with one widely rounded apex ” (Nabozhenko & Steiner 2021: 571). This character can be more precisely defined as the margin of the ostium of the penis. The penis of Tenebrionidae is a tube-like structure which is supported by a Y- or wishbone-shaped longitudinal sclerotized portion called the penis rod (Doyen 1966) or median-lobe baculus (Matthews & Bouchard 2008). At the distal end of the penis, between the two forks of the penis rod or baculus, is the ostium, or the opening through which the internal sac is everted during copulation (for definitions and drawings, see Lawrence & Ślipiński 2013 and Doyen 1966). Thus, the character discussed here is reporting that the ostium of Doyenellus is bounded above by a single rounded margin while ventrally it is incised medially. All New World Helopina we have examined have the ostium located apicoventrally where the opening can be difficult to characterize depending on the preservation and state of the internal sac which is often slightly protruding and may obscure the true boundary of the ostium. The ostium of Doyenellus (Fig. 3 C) is accurately described by Nabozhenko & Steiner (2021) but the variation seen across New World Helopina does not allow for simple attribution of character states across species. For instance, some species have a much more deeply incised ventral margin of the ostium (Fig. 3 D) while others seem to have a fully bilobed apex of the penis where both the top and bottom are incised (Fig 3 E). This seems to be an important character system that could diagnose particular species but we see no consistent or easily binary character states which separate phylogenetically distinct genera. The incised ventral margin of the ostium is clearly not unique to Doyenellus. 5. Female genital tube with the “ presence of one-way valve (valve without short processes laterally) in base of accessory gland duct (unique character within American Helopini) ” (Nabozhenko & Steiner 2021: 571). Nabozhenko & Steiner (2021: 572) state that this character is “ unusual within American Helopini but not unique within Helopini of the world. ” We only examined the internal female genital tube of two species (the robust flightless species Helops opacus LeConte, 1859 from California, and an undescribed narrow flighted species from Arizona which is very consistent with Central American Tarpela). Both had all the same characteristics as described for Doyenellus and possessed a one-way valve at the base of the accessory gland (Fig. 3 G). Nabozhenko & Steiner (2021) did not mention how many other New World species they may have examined, but this state is not unique and our limited sampling suggests it is likely common if not ubiquitous. 6. Leg armature characterized by “ dilated male pro- and mesotarsi, very long setae around the tibial apex ” (Nabozhenko & Steiner 2021: 571). This character was included in the structures which putatively diagnose Doyenellus but was not reported as unique to the genus, which it is not. Dilated male tarsi may be more common among flightless species, but we have observed this across all of the other genera currently recognized for North American Helopina. All six of the character systems presented as diagnostic for Doyenellus are inadequate to readily diagnose a genus among the North American Helopina. The three species currently placed in the genus may be closely related, with D. cisteloides being particularly apomorphic, especially in the secondary sexual characters discussed above, and the two other species presenting, at best, variations on that theme. However, these putatively diagnostic characters were not satisfactorily surveyed across the broader fauna during the original description and they do not serve to adequately diagnose a separate genus.	en	Aalbu, Rolf L., Johnston, M. Andrew (2025): Reorganization of New World Helopini (Coleoptera: Tenebrionidae: Tenebrioninae). Zootaxa 5631 (1): 103-120, DOI: 10.11646/zootaxa.5631.1.4, URL: https://doi.org/10.11646/zootaxa.5631.1.4
795187DDFFC9EF0EF4D5D24EEFF1E7D7.taxon	description	As discussed above, Nautes and Tarpela are together separated from “ Helops ” primarily on the basis of flight. This is not a binary system, however. Some species, like Helops arizonensis Horn, 1874 have fully macropterous and flighted males while females have wings shorter than the elytra and have not been observed to fly to lights where males are readily collected. Indeed, even for species of “ Helops ” which have historically been classified as flightless, largely due to the distance between the middle and hind coxae and the general body form, possess reduced metathoracic wings (Fig. 4). Nautes, Tarpela as well New World “ Helops ” have been found to be without any dependable genus-level defining morphological character (s) and thus not satisfactorily distinguished in any keys. This has discouraged taxonomic works on North American helopines due to the inability of new species to be reliably placed to genus. Nabozhenko & Ando (2018: 283) agreed and suggested that “ The main and indistinct differences between Nautes-Tarpela and New World Helops are in the structure of the prosternal process and mesoventrite. Tarpela and Nautes have projected prosternal process and very deeply depressed mesoventrite with strongly elevated and flat or weakly convex lateral margins of mesocoxal process. ” This is true of the type species of both Tarpela and Nautes but does not fit into the huge amount of variation we found in both the prosternal process and the mesosternal excavation in the 147 species we examined. We outline these variations below: Prosternal process: 1. completely declivious rounded behind procoxae. (Fig. 8) 2. straight but declivious (various angles) (Fig. 7) 3. declivious rounded then straight 4. declivious straight then upcurved (Fig. 9) 5. completely straight (Fig. 11) Mesosternal lateral elevations and mid cavity: 1. completely flat (Fig. 7) 2. slightly inflated, cavity shallow (Fig. 8) 3. slightly inflated, cavity deep 4. moderately inflated, cavity shallow 5. moderately inflated, cavity deep (Fig. 9) 6. strongly inflated, rounded, cavity shallow 7. strongly inflated, rounded, cavity deep (Fig. 10) 8. strongly inflated, angulate, cavity deep (Fig. 11) Various combinations of these characters exist and some variation within species was also found. Prosternal processes vary from completely declivious to declivious but then straight to then upcurved to partially declivious. Mesosterna vary from completely flat to slightly inflated to moderately inflated to strongly inflated but rounded (as in both Nautes fervidus Pascoe and Tarpela browni Bates) to strongly inflated with anterior apex angulate as in T. catenata Champion, T. costata Champion, T. thoracica Champion (Fig. 12) and T. recticulata Champion. However, T. occidentalis (Allard) from Jamaica (Fig. 13) which would seem to fall in this group morphologically, the prosternal process is decumbent to decumbent and sometimes upcurved and the mesosternum is shallow with sides only slightly inflated. The inconsistency and wide variation of both the prosternal process and the mesosternum, even within a single species discounts using these characters as generic differential characters.	en	Aalbu, Rolf L., Johnston, M. Andrew (2025): Reorganization of New World Helopini (Coleoptera: Tenebrionidae: Tenebrioninae). Zootaxa 5631 (1): 103-120, DOI: 10.11646/zootaxa.5631.1.4, URL: https://doi.org/10.11646/zootaxa.5631.1.4
795187DDFFC5EF0CF4D5D3C3E8ADE7DF.taxon	type_taxon	Type species: Dinomus perforatus Brême, by monotypy.	en	Aalbu, Rolf L., Johnston, M. Andrew (2025): Reorganization of New World Helopini (Coleoptera: Tenebrionidae: Tenebrioninae). Zootaxa 5631 (1): 103-120, DOI: 10.11646/zootaxa.5631.1.4, URL: https://doi.org/10.11646/zootaxa.5631.1.4
795187DDFFC5EF0CF4D5D3C3E8ADE7DF.taxon	diagnosis	Diagnosis. Body variable, 4 - 20 mm long, shining and submetallic to matte and piceus, often shining brownish black. Eyes reniform, facets small, longitudinally compressed in dorsal view, with or without marginal groove; gena without longitudinal groove beneath eye; antennae with 11 antennomeres, variable, terminal antennomere rounded or elongate, always asymmetrical. Femora without spines; tarsi often with yellow silky setae beneath, but ranging from plantar surface lined with stout black setae to large flattened tomentose pads. Epipleuron ending near apex of fourth ventrite, not attaining elytral apex. Male aedeagus of the catomoid form, with parameres much shorter than phallobase, with apical two thirds covered in short anteriorly directed spinose setae.	en	Aalbu, Rolf L., Johnston, M. Andrew (2025): Reorganization of New World Helopini (Coleoptera: Tenebrionidae: Tenebrioninae). Zootaxa 5631 (1): 103-120, DOI: 10.11646/zootaxa.5631.1.4, URL: https://doi.org/10.11646/zootaxa.5631.1.4
795187DDFFC5EF0CF4D5D3C3E8ADE7DF.taxon	discussion	Remarks. The synonymies presented for this genus follow from the reassessment of the genera presented above. We believe that flight has been lost multiple times (and can even be sexually dimorphic such as in Dinomus arizonensis) in this lineage and that corresponding differences in body convexity, elytral shape, pro- and mesosternal morphology are a result. Moreover, our expanded unified genus concept, which follows that proposed by Horn (1870), more truly aligns with available phylogenetic data and makes this group readily diagnosable.	en	Aalbu, Rolf L., Johnston, M. Andrew (2025): Reorganization of New World Helopini (Coleoptera: Tenebrionidae: Tenebrioninae). Zootaxa 5631 (1): 103-120, DOI: 10.11646/zootaxa.5631.1.4, URL: https://doi.org/10.11646/zootaxa.5631.1.4
795187DDFFC5EF0CF4D5D3C3E8ADE7DF.taxon	distribution	Table 1 outlines the proposed nomenclatural changes in New World Helopini along with the current name and species examined. Full nomenclatural and bibliographic data can be found in Bousquet et al. (2018) and Bouchard et al. (2021). These nomenclatural changes necessitate renaming one species: Dinomus horni Aalbu & Johnston nom. nov. for Helops perforatus Horn, 1880: 153. (Fig. 13) Distribution: USA: Texas. Secondary homonym of Dinomus perforatus Brême, 1842: 24.	en	Aalbu, Rolf L., Johnston, M. Andrew (2025): Reorganization of New World Helopini (Coleoptera: Tenebrionidae: Tenebrioninae). Zootaxa 5631 (1): 103-120, DOI: 10.11646/zootaxa.5631.1.4, URL: https://doi.org/10.11646/zootaxa.5631.1.4
