Rudolfina Rohacek 1987

Genus Rudolfina Roháček, 1987 Rudolfia Roháček, 1982: 225 (feminine, a junior homonym of Rudolfia Wilson, 1924 (Crustacea: Copepoda); type species: Limosina rozkosnyi Roháček, 1975, by monotypy). Rudolfina Roháček, 1987: 474 (feminine, nom. nov. for Rudolfia Roháček, 1982; type species: Limosina rozkosnyi Roháček, 1975, automatic). Rudolfia – Roháček 1982: 225 [formal diagnosis, phylogeny]; 1983: 152–154 [full description, phylogenetic notes]; 1987: 474 [homonymy]. — Marshall & Richards 1987: 999–1000 [diagnosis in key, illustr.]. Rudolfina – Roháček 1998: 483 [diagnosis, illustration]. — Marshall & Buck 2010: 1180–1183 [diagnosis in key, phylogenetic notes, biology]. — Su et al. 2017: 392–393 [key]. Redescription BODY. Colour light to dark brown. Length 1.4–2.3 mm. HEAD. With 3–5 interfrontal setae (of equal length or the foremost shorter), 1 (rarely 2) inclinate orbital setula and 4–10 small orbital setulae inside and below 2 strong, exclinate orbital setae; ocellar triangle with pair of strong setae and 3–5 additional small setulae; outer vertical seta strong, exclinate; inner vertical seta inclinate; occipital and paravertical setae inclinate, well-developed; postocellar seta inclinate, weakly developed. Eye-to-gena height ratio variable between species (1.5:1 to 3.5:1). Vibrissa strong. Gena with 1–2 strong subvibrissal setae and 4–9 smaller setulae. THORAX. Surface pruinose. Postpronotal lobe with 2–3 setae, outer seta strong, inner seta(e) reduced. Notopleural seta, 2 supra-alar setae and prescutellar dorsocentral seta strong. Acrostichal setulae in 4–8 rows, with 1 enlarged prescutellar acrostichal seta (almost as long as dorsocentral). Katepisternum with strong elongate posterior seta and reduced anterior seta. LEGS. Fore femur with 3–5 elongate setae dorsally (except R. exuberata sp. nov.). Fore tibia with 3–5 elongate setae ventrally. Mid femur with row of 3–10 anterodorsal setae extending from base, row of 2–5 dorsal setae on apical ¼, and basal cluster of 4–21 small setae ventrally; males usually with additional ventral seta (often in ventrobasal cluster). Mid tibia with 4 dorsal setae (basal anterodorsal, medial anterodorsal distal anterodorsal and distal posterodorsal); males with ventral comb of 4–13 setae on apical ½ or less (, R. exuberata sp. nov. and R. remiforma sp. nov. with setae of ventral comb weakly developed); females usually with midventral seta (absent in R. megepandria sp. nov.). Hind tibia with small apical ventral spur. WING. Always fully developed, with wing tip reaching or exceeding apex of abdomen. Costa extending to or just beyond end of R 4+5, and with single costagial seta> 2.0 × length of nearby setae. R 4+5 slightly curved towards costa distally. Cell dm with short stub veins of M 1 and Cu-A1 extending beyond dm-cu. Alula narrow, posterior margin straight. ABDOMEN. Sternites and tergites well sclerotized and setose (posterior and lateral margins more densely setose). Male sternite 4 usually simple (rarely densely setose medially). MALE ABDOMEN. Posterior margin of sternite 5 with lobe on each side of medial emargination (shape and size of emargination and lobes vary among species). Transverse (ventral) portion of sternite 6 narrow; straight or weakly arcuate. Ring sclerite (in the right membrane of segment 7, possibly derived from a spiracle) large and distinct. Epandrium setose, often with larger setae lateral to anal opening, and with right anteroventral corner drawn out into a finger-like process that extends to the hypandrium. Male cercus usually distinct, fused with the epandrium (reduced and obscured beneath the epandrium in a few species; e.g., R. pilosa sp. nov., R. remiforma sp. nov.). Hypandrium (Fig. 3B) Y-shaped with emarginate posteromedial extension; hypandrial arms posteriorly deeply bilobed posteriorly, with lateral lobe articulating with the epandrium and the medial lobe articulating with posterolateral corner of hypandrium. Pregonite distinct, small, near anterior base of postgonite. Postgonite generally simple and slender, with 3–4 setulae on anterior margin but modified in some species; ejaculatory apodeme small and finger-like, with small globular sperm pump, usually close to the basiphallus (easily lost during dissection); basiphallus simple (without an epiphallus); distiphallus with distinct elongate dorsal sclerite; acrophallus with dorsolateral lobes and a single ventral sac (often reduced). FEMALE ABDOMEN. Tergite 8 apparently tripartite, with two lateral triangular sclerites and a medial sclerite (reduced in several species). Epiproct bare except for usual pair of small setae and a few scattered setulae, strongly sclerotized, and fused laterally with cerci (except in R. cavernicola). Cercus with single flattened apical seta. Sternite 7 variable. Sternite 8 weak, transverse, covered in small setulae; pair of small, bispinose plates along posterior margin. Hypoproct very narrow, forming horseshoe-shaped band immediately below the cerci. Spermathecae (1 pair + 1 single) generally disc-shaped or lenticular, with thin, long sclerotized ducts. Related and similar genera All species of Rudolfina will key out to “ Rudolfia ” in the key to Nearctic Sphaeroceridae by Marshall & Richards (1987) but they will key out as “ Archiceroptera genus complex, in part” at couplet 72 in the key of Marshall & Buck (2010) to Neotropical Sphaeroceridae. This previous treatment reflected uncertainty about the limits between Rudolfina and the many undescribed Neotropical species in the Archiceroptera genus complex. The Archiceroptera complex is part of a larger group of Limosininae (including Aptilotella Duda, 1924, Archiceroptera Papp, 1977, Bitheca Marshall, 1987, Bromeoloecia Spuler, 1924, Pterogramma Spuler, 1924 and Robustagramma Marshall & Cui, 2005) characterized by an unusual process extending medially from the lower right margin of the epandrium to the hypandrium. The relationships within this group need further study but the morphological analysis by Paiero (2017) suggests that Rudolfina is closely related to Bromeloecia. Within this group, Rudolfina resembles Archiceroptera in characters of the female epiproct and cercus. However, in Archiceroptera the epiproct is completely desclerotized medially (anteriorly sclerotized in Rudolfina), the cercus is separate from the epiproct and has a partially concave inner margin, and sternite 8 is divided into a pair of elongate lateral sclerites without the paired setulose sclerites found in Rudolfina. Archiceroptera species also differ from Rudolfina in having M 1 extending as a pseudovein to the wing margin, CuA 1 rarely with a distinct stub vein, the male cercus free from the epandrium and with a distinct ventral process, and (in many, but not all species) five or more dorsal mid tibial setae and two or more inclinate orbital setulae. Biology Roháček (1987) recorded R. rozkosnyi from dung and occasionally from mud and decaying vegetation, but most of the new species considered here were collected in dung or carrion traps. Larvae remain unknown. Distribution Rudolfina has a mostly western Nearctic montane distribution, with high endemism in the southwest and the mountains of Mexico (Sierra Madre del Sur, Sierra Madre Oriental and Sierra Madre de Chiapas). Two widely separated species occur in the Palaearctic region (R. rozkosnyi and R. zhangi) and one species (R. exuberata sp. nov.) is widespread at low elevations from the southern United States to South America. Other than R. exuberata sp. nov., no true species of Rudolfina are known from south of Guatemala. Other Neotropical species previously treated as Rudolfina are discussed below. Results of the phylogenetic analysis Twenty-seven most parsimonious trees were generated, summarized here as a strict consensus tree (Fig. 5) and a majority rules consensus tree (Fig. 6). Characters were optimized on one of the equal length trees (Fig. 7) which was selected based on the recovery of several groups supported by putatively higher weight characters. Shared male genitalic morphology supported a close relationship of Rudolfina bucki sp nov. with R. megepandria sp. nov. and R. tumida sp. nov. with [R. rozkosnyi + R. digitata + R. zhangi]. The combined elongation of the epiproct and female cercus suggests that R. newtoni sp. nov. is closely related to [R. exuberata sp. nov. + R. pauca sp. nov. + R. remiforma sp. nov.], although the form of the epiproct in R. newtoni sp. nov. is apparently intermediate between the strongly elongated form found in the R. exuberata clade and the shorter epiproct of other Rudolfina. All trees recovered R. cavernicola as a sister taxon to the remaining species, which form a monophyletic group characterized by the fusion of the female cercus with the posterolateral corner of the epiproct, the elongation of the medial part of tergite 8 and by characters of the male cercus and surstylus. This tree suggests a New World origin for Rudolfina. Within Rudolfina excluding R. cavernicola, R. rozkosnyi, R. digitata and R. tumida sp. nov. appear to be a basal grade predating the origin of a clade comprising the Mexican-Guatemalan species. These four species all have a laminate lobe on the surstylus, apparently derived from the simple laminate margin of R. cavernicola (absent in other species). The largely Mexican-Guatemalan clade can be recognized by the simple, rounded anterior lobe of the male surstylus and the absence of dorsal swellings on the dorsal sclerite of the distiphallus. The R. exuberata group (including R. exuberata sp. nov., R. remiforma sp. nov. and R. pauca sp. nov.) is characterized by a small elongate male cercus, tulip-shaped epiproct, and reduction of the female cercus. Rudolfina remiforma sp. nov. and R. pauca sp. nov. are known from only a few localities at higher elevations, as is typical of the genus, but the widespread R. exuberata sp. nov. occurs at much lower elevations than its more localized congeners. Key to the New World Rudolfin

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