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  Beetles that live with ants ( Carabidae, Pseudomorphini, Pseudomorpha   Kirby, 1825  ) ... 29 Beetles that live with ants (Carabidae, Pseudomorphini, Pseudomorpha  Kirby, 1825): A revision of the santarita  species group erry L. Erwin 1,† , Lauren M. Amundson 2,‡ 1  Hyper-diversity Group, Department of Entomology, MRC-187, National Museum of Natural History, Smithsonian Institution, Washington, P.O. Box 37012, DC 20013-7012, USA 2  Hyper-diversity Group Summer Intern, Department of Entomology, MRC-187, National Museum of Natural History, Smithsonian Institution, Washington, P.O. Box 37012, DC 20013-7012, USA †  ‡  Corresponding author:   Terry L. Erwin  ( Academic editor:   L. Penev   | Received 24 September 2013 | Accepted 24 November 2013 | Published 13 December 2013 Citation:  Erwin L, Amundson LM (2013) Beetles that live with ants (Carabidae, Pseudomorphini, Pseudomorpha   Kirby, 1825): A revision of the santarita   species group. ZooKeys 362: 29–54. doi: 10.3897/zookeys.362.6300 Abstract Te Western Hemisphere genus Pseudomorpha   Kirby 1825 was last revised by Notman in 1925 based on only a few known species (22) and paltry few specimens (73); other authors have added an additional six species represented by 53 additional specimens since 1925. Baehr (1997) assigned three species from  Australia to this genus, albeit in a new subgenus,  Austropseudomorpha   Baehr 1997. A recent study of col-lections from throughout the Americas (1757 specimens) has revealed numerous new species that can be arrayed across 19 species groups based on a suite of attributes, some used by Notman and others newly discovered. A taxonomic revision of the species contained in one of these species groups, santarita  , is pro-vided herein, as well as a distributional synopsis of the remaining 18 species groups. New species described herein are as follows, each with its type locality: Pseudomorpha huachinera    sp. n. , Arroyo El Cocono, Sierra Huachinera, Sonora, México;   P.    patagonia    sp. n. , Madera Canyon, Santa Rita Mountains, Arizona; P.    penablanca    sp. n. , Peña Blanca Lake, Arizona; P. pima    sp. n. , Madera Canyon (lower), Santa Rita Moun-tains, Arizona; P. santacruz    sp. n. , Madera Canyon, Santa Rita Mountains, Arizona; and P. santarita    sp. n. , Santa Rita Ranch, Santa Rita Mountains, Arizona.   ZooKeys 362: 29–54 (2013)doi: 10.3897/ Copyright Terry L. Erwin, Lauren M. Amundson. This is an open access article distributed under the terms of the Creative Commons Attribution International License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the srcinal author and source are credited. RESEARCH ARTICLE Launched to accelerate biodiversity research A peer-reviewed open-access journal  Terry L. Erwin & Lauren M. Amundson / ZooKeys 362: 29–54 (2013) 30 Resumen El género Pseudomorpha   Kirby 1825 del hemisferio occidental fue revisado la última vez por Notman en 1925 basado en solo unas pocas especies conocidas (22) y pocos especímenes muy pobres (73); desde 1925 otros autores añadieron 6 especies representadas por 53 especímenes adicionales. Baehr, en 1997, asignó tres especies de Australia a este género, aunque en un subgénero nuevo,  Austropseudomorpha   Baehr 1997. Un estudio reciente de las colecciones a través de las Américas (1757 especímenes) ha revelado numerosas especies nuevas que pueden ser ordenadas en 19 grupos de especies basadas en una serie de atributos, algunos usados por Notman y otros recién descubiertos. Una revisión taxonómica de las espe-cies contenidas en uno de estos grupos de especies, santarita  , se provee aquí, así como una sinopsis dis-tribucional de los otros 18 grupos. Las nuevas especies descritas aquí son las siguientes, cada una con su localidad tipo: Pseudomorpha huachinera sp. n. , Arroyo El Cocono, Sierra Huachinera, Sonora, México; P. patagonia    sp. n. , Madera Canyon, Santa Rita Mountains, Arizona; P. penablanca    sp. n. , Peña Blanca Lake, Arizona; P. pima    sp. n. , Madera Canyon (bajo), Santa Rita Mountains, Arizona; P. santacruz  , sp. n. , Madera Canyon, Santa Rita Mountains, Arizona; and P. santarita   s p. n. , Santa Rita Ranch, Santa Rita Mountains, Arizona. Keywords False-form beetles, new species, new species groups, identification key, distributions, male genitalia, fe-male ovipositor, Hymenoptera: Formicidae Palabras clave Carábidos atípicos, nuevas especies, nuevos grupos de especies, clave de identificación, distribuciones, genitalia del macho, ovipositor de la hembra, Hymenoptera: Formicidae Introduction Species of Pseudomorpha   and some related genera are obligatory myrmecophiles in their larval stages. All known species of Pseudomorpha   are terrestrial and their ant hosts live in the soil; however, adults of the pseudomorphine genus Samiriamorpha   have been found in the arboreal nests of  Azteca   ants (Erwin and Geraci 2008). Te genus Pseudomorpha   currently includes 27 described species in the Western Hemisphere and more than 125 undescribed species (Erwin in prep.). Te present contribution pro-vides information on six undescribed species in the new santarita   species groups with a key for identification of those species. Notman (1925) did not have specimens of this group; therefore, they do not run to any couplet he provided in his key. With the cur-rent treatment, we start bringing organization to one of the last poorly known carabid lineages in North America particularly, but also in Middle and South America. Tis information will begin to aid those interested in ants and their commensals, as well as collection managers in understanding what is present in their collections and how it is to be ordered.Erwin and Geraci (2008) provided a complete history of work on this group of carabid beetles in the Western Hemisphere; Baehr (1992, 1997) did the same for the  Beetles that live with ants ( Carabidae, Pseudomorphini, Pseudomorpha   Kirby, 1825  ) ... 31 Eastern Hemisphere species. Te only monograph published of the Western Hemi-sphere species is that of Notman (1925). His key to the species known to him at the time, as it turns out, is more of a key to species groups now that we know there are over 125 species in Pseudomorpha  , alone. Lenko (1972) and Erwin (1981) described larvae and Liebherr and Kavanaugh (1985) noted that species of Pseudomorpha   are ovovi-viparous. Since Erwin and Geraci (2008), an additional new genus was discovered in Guyane (French Guiana) (Erwin 2013) Specimens and methods Included in the overall study of this genus are a total of 1757 specimens from the National Museum of Natural History, Washington, DC (NMNH, erry L. Erwin, Curator) and several other institutions and private collections (see Appendix 1 for the specimens covered in the current paper).Methods and species concepts follow those previously described (Ball 1959; Er-win and Kavanaugh 1981; Kavanaugh and Erwin 1991). Te species validation and diagnosis format follows as closely as possible that suggested in Erwin and Johnson (2000). Measurements of length (ABL, SBL) and width (W) follow those of Ball (1972) and Kavanaugh (1979): ABL (apparent body length), measured from apex of labrum to apex of the abdomen; 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 (pronotal length ), measured from apical to basal margin along midline, and LE (elytron length), measured from apex of scutel-lum to apex of the longer elytron; and W (total width), measured across both elytra at their widest point with suture closed. Measures and ratios are presented in the tables in Appendix 2.Habitus and attribute images of the adult beetles portray most of the character states referred to in the key provided. Male and female genitalic presentations are standard for descriptive taxonomy of carabid beetles, and in this case are digital photo-illustrations (Erwin 2011). Te images of the adult and its parts were made with a Visionary Digital M  high resolution imaging system. Figure captions include an ADP number, which is a unique identification number for the specimen that was illustrated or imaged and links the specimen and associated illustrations and/or image to addi-tional information in electronic databases at the NMNH.Geographical data are presented based on all known specimens of each species available at the time of manuscript preparation. Georeferences have been determined from locality information provided on specimen labels. Latitude and longitude are reported in decimal degrees. A distribution map is provided for the species (Fig. 18). Herein, an English vernacular name is proposed, as vernacular names are becoming increasingly needed in conservation and/or agricultural and forestry applications, as well as for the Encyclopedia of Life ( ).  Terry L. Erwin & Lauren M. Amundson / ZooKeys 362: 29–54 (2013) 32 Accounts of taxa Pseudomorphini Newman, 1842 beetlesPseudomorphini Newman, 1842:365 (as Pseudomorphites) axonomy. Stable at the generic level. Classification.  According to Ober and Maddison (2008), Pseudomorphini appears as a branch of the higher Carabidae and associated with Graphipterini and Orthogonini; according to Erwin and Geraci (2008), the adelphotaxon is the tribe Orthogonini. All three tribes are associated in some way with ants or termites. Male genitalia of pseudo-morphines have a bonnet-shaped phallobase like the lebiomorphs, yet their accompany-ing parameres are large and nearly symmetrical (and in some species the parameres are sparsely setiferous, as in some primitive lineages of the family). Many known lineages of Pseudomorphini have been so highly selected for life with ants (and possibly termites) that external structures do not help much in discovering more normal carabid relatives. References. Baehr (1992, 1997); Erwin and Geraci (2008); Moore (1964, 1974, 1983); Ogueta (1967); Notman (1925). Pseudomorpha   Kirby, 1825  Pseudomorpha   Kirby, 1825:98 Heteromorpha   Kirby, 1825:109  Axinophorus   Dejean, 1829:174 Drepanus   Dejean, 1831:434 Heteromorphus   Chaudoir, 1852:63 ype species. Pseudomorpha excrucians   Kirby, 1825:101 Proposed English vernacular name.  Western False-form beetles Number of described Western Hemisphere species. 27 Current known number of undescribed Western Hemisphere species. 125 Number of described Australian species. 3 (Subgenus  Austropseudomorpha   Baehr, 1997)  Adelphotaxon. Tuxtlmorpha   Erwin & Geraci, 2008 + Samiriamorpha   Erwin & Geraci, 2008 (see Erwin and Geraci 2008 for phylogeny). axonomy. Stable at the generic level (Erwin 2013; Erwin and Geraci 2008), although many undescribed species need to be treated (Erwin in prep.). All of these undescribed species have been assigned to the species groups listed herein and their male genitalia have been illustrated, their label data entered into a database, and their localities mapped using Google Earth Pro.  Beetles that live with ants ( Carabidae, Pseudomorphini, Pseudomorpha   Kirby, 1825  ) ... 33 Diagnosis. Form moderately depressed or rarely subcylindrical, narrow or broad, lean or robust, head visible from above, legs concealed beneath when in repose. Color ranges from black to light brown, rarely slightly rufous; only adults of P. excrucians   Kirby, Yasuni-morpha piranhna   Erwin & Geraci, Guyanemorpha spectablis   Erwin from Guyane are mark-edly bicolored. Head with mouthparts visible in dorsal aspect; ventrally beneath eye with deeply recessed groove for insertion of antennal base; mandibular scrobe nearly effaced, delimited by row of short stout setae; mentum and submentum fused; antennal scape par-tially visible in dorsal aspect. Anterior coxal cavities closed, median coxal cavities conjunct, metepimeron visible. Abdomen with six visible sterna, sternum III with broad medial emar-gination on posterior margin; sterna V and VI in male with dense row of decumbent and yellowish robust setae medially. Male parameres long, nearly of same length (more or less symmetrical), glabrous or setose, not balteate; phallobase bonnet-shaped, crested or not.  Way of life.  As far as is known, adults are found in and around ant nests and in the surrounding vicinity; females are ovoviviparous (Liebherr and Kavanaugh 1985); larvae are known to be ant nest inquilines (Lenko 1972; Erwin 1981), or perhaps liv-ing with termites (Ogueta 1967). Of adults found at lights (UV, MV, and white light), most are males. Geographic distribution. Members of this genus are known to occur from Ore-gon, Idaho, and Colorado in the north to Argentina in the south, including the Carib-bean area, and in southern Australia. Tey should be looked for in southern Wyoming, where they are also likely to occur. Te only eastern species, P. excrucians   Kirby, is related to species from the Caribbean and South America, not to those lineages from the American west and southwest. Habitat. Dry loamy or sandy soil where ants prefer to build nests from MASL -72m to 2606m altitude in deserts, grasslands, and open and closed forests. Description. Head   (cf. Figs 7–12) with two supraorbital setigerous punctures per eye near their posterior corner, however, numerous accessory setae in some groups obscure them; frontal impressions absent. Clypeus markedly wide, trapezoidal with rounded an-terior angles and shallowly lobed posterior margin; posterior margin in some adults very shallow, or effaced, bearing a single long seta each side near anterior corner. Eyes flat, or slightly convex; small gena with numerous stout setae. Antenna of varying length, either shorter, or longer than distance from antennal base to anterior coxae; antennomeres 3–9 slightly wider distally and appearing flattened. Labrum visible, about 2/3 as wide as cl-ypeus, rectangulate, bearing six setae along anterior margin. Mandible markedly flattened with a very short and acute apex; outer margin ventral of the scrobe with short stout setae. Maxillary palpi markedly short, 3-segmented, palpomeres slightly depressed, palpomere 3 truncate apically. Labial palpus with short bisetose palpomere 2; palpomere 3 markedly securiform and robust, its distal margin mostly membranous with sensory organs. Prothorax.  Pronotum (cf. Figs 1–6) wider than head, transverse, with broadly explanate margins, or in cylindrical species narrowly explanate margins; without a pair of setigerous punctures each side, apical, lateral and posterior margins with border of stout setae; hind angles obtuse, broadly rounded. Proepisternum with prosternal pro-cess multisetiferous apically, intercoxal process feebly margined.
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