Descriptions of four new species of Minyomerus Horn, 1876 sec. Jansen & Franz, 2018 (Coleoptera: Curculionidae), with notes on their distribution and phylogeny

This contribution adopts the taxonomic concept approach, including the use of taxonomic concept labels (name sec. [according to] source) and region connection calculus-5 (RCC–5) articulations and alignments. Prior to this study, the broad-nosed weevil genus Minyomerus Horn, 1876 sec. Jansen & Franz, 2015 (Curculionidae [non-focal]: Entiminae [non-focal]: Tanymecini [non-focal]) contained 17 species distributed throughout the desert and plains regions of North America. In this review of Minyomerus sec. Jansen & Franz, 2018, we describe the following four species as new to science: Minyomerus ampullaceus sec. Jansen & Franz, 2018 (henceforth: [JF2018]), new species, Minyomerus franko [JF2018], new species, Minyomerus sculptilis [JF2018], new species, and Minyomerus tylotos [JF2018], new species. The four new species are added to, and integrated with, the preceding revision, and an updated key and phylogeny of Minyomerus [JF2018] are presented. A cladistic analysis using 52 morphological characters of 26 terminal taxa (5/21 outgroup/ingroup) yielded a single most-parsimonious cladogram (Length = 99 steps, consistency index = 60, retention index = 80). The analysis reaffirms the monophyly of Minyomerus [JF2018] with eight unreversed synapomorphies. The species-group placements, possible biogeographic origins, and natural history of the new species are discussed in detail.


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This phylogenetic study follows Jansen & Franz (2015) in the use of the taxonomic concept approach; see characters. This manuscript is arranged with the species descriptions appearing first, followed by the key 150 to species, and then by the phylogenetic and RCC-5 alignment results.

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Phylogenetic analysis 152 The morphological cladistic analysis includes 26 terminal taxa; with 21 ingroup and 5 outgroup terminals. 153 The ingroup terminals were represented by 17 species previously assigned to Minyomerus  154 and four newly recognized species. In keeping with our previous analysis, we sampled outgroups fairly  The character matrix was edited and phylogenetic results viewed using the WinDada and WinClados 168 interfaces of WinClada, respectively (Nixon et al. 2002). Characters are numbered in accordance with 169 descriptive sequence used in the species accounts. A "-" symbol indicates inapplicable (character, state), 170 whereas a "?" symbol indicates missing information, e.g., due to the unavailability of male specimens or 171 insufficient specimens on hand to permit full dissections. Characters 9, 27, 39, 45 -47, 49, and 51 were 172 mapped onto the preferred phylogeny using ACCTRAN optimization (see Agnarsson & Miller 2008), and 173 the remaining characters had an unambiguous optimization. All multi-state characters but one were coded 174 as additive, as explained beneath the description for each character (see Phylogenetic Results), based 175 on their alignment with the preferred phylogeny. Each alternative coding scheme was tested both alone 176 and in unison with the other multi-state characters to assess their impact on the topology of the preferred 177 phylogeny. 178 The most parsimonious tree and character state optimizations were inferred under parsimony using 179 NONA (Goloboff 1999). An unconstrained heuristic search was conducted using the commands: hold 180 100001, mult * 1000, hold/100, with mult * max * selected. Bootstrap support was inferred in 181 WinClada using the parameters of 1000 replications, hold 1000, hold/100, mult * 10, 182 Don't do max * , and Save consensus. Finally, Bremer support values (Bremer et al. 1994) and 183 relative fit difference (Goloboff & Farris 2001) were calculated in NONA using the commands: hold 184 1001, sub 20, bs for Bremer support values, and bs * for relative fit difference, respectively (Goloboff 185 et al. 2008). 186 The motivation for providing Bremer support values and relative fit difference comes from their respective interpretations, based on how the measures are calculated, per Goloboff & Farris (2001). Both of these indices rely on summation of the number of favorable and contradictory characters when comparing a most-parsimonious tree to a suboptimal tree. If the step length of the i th character (I) of n total characters on the most-parsimonious tree (L MPT ) is less than its corresponding step length on the suboptimal tree (L SUB ), the character is designated as favorable ( f i ), but if the opposite is true, the character is designated as contradictory (c i ), and expressed formally:

Manuscript to be reviewed
Where the number of favorable (F) and contradictory (C) characters are defined, respectively, as: Bremer support values (bsv) and relative fit difference (rfd) are then calculated simply as: The Bremer support value for a node thus indicates how many more characters support a node than         tooth, which is approximately equilateral and whose sides are sub-equal in length to surrounding setae.

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Elytra with 10 complete striae; striae shallow; punctures faint beneath appressed scales, separated by   Named in reference to the long, somewhat unkempt, erect setae on the anterior margin of the pronotum-445 franko = "free"; Old High-German adjective (Brown 1956).   The indication of "Sphaeralcea hastula A. Gray" is not a valid name and appears to be a misspelling of   Manuscript to be reviewed Elytra Length/width ratio 3.12-3.16; widest at anterior 1/5; anterior margins jointly 1.5-2× wider 514 than posterior margin of pronotum; lateral margins gently converging after anterior 1/5, more strongly 515 converging in posterior 1/4. Posterior declivity angled at 65-70°to main body axis. Elytra with 10 516 complete striae; striae broadly sculpted; punctures faint beneath appressed scales, separated by 5-7× their 517 diameter; intervals elevated, with every second interval, beginning at elytral suture, more strongly raised 518 than adjacent intervals.

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Abdominal sterna Ventrite III anteromesally incurved around a fovea located mesally on anterior 520 margin, posterior margin elevated and set off from IV along lateral 1/3s of its length. Sternum VII mesally 521 2/3× as long as wide; anterior margin straight.

Distribution
This species has been found in three localities along the Snake River in Idaho (USA), and is thought to be endemic to the Snake River Plain (Fig. 38). sub-recumbent to sub-erect, tan to brown in color.

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Mandibles Covered with white scales, with 2-3 longer setae, and 1-3 shorter setae between these.   Named in reference to the short, apically explanate setae interspersed throughout the dorsum, which give 666 this species a distinctly "knobbed" appearance; tylotos -knobby; Greek adjective (Brown 1956).   Body shape distinctly flask-like, with strongly constricted, sub-cylindrical pronotum and greatly protuberant elytra; in dorsal view, elytra nearly 2× width of pronotum at maximum width and nearly 3/4× as wide as long; in lateral view, anterior and posterior declivities of elytra abrupt and nearly vertical, with anterior elytral margin projecting strongly and characteristically dorsad of articulation with posterior pronotal margin; corpus of spermatheca with highly elongate projection aligned with midline of the ramus, which is basally tapered and angled at nearly 45°to corpus . Manuscript to be reviewed -Body shape usually narrow; elytra typically not more than 1.5× width of pronotum and typically not more than 2/3× as wide as long in dorsal view; elytral declivities in lateral view variable, but anterior margin never abruptly and strongly projected dorsad of posterior pronotal margin; spermatheca variable, but never with elongate projection aligned with midline of ramus . Anterior margin of pronotum bearing strikingly long setae, which project laterally up to 80°from longitudinal body axis and at least equal to diameter of eye; spermatheca with short, somewhat bulbous corpus, ramus sub-equal in size and perpendicular to corpus, and collum strongly recurved along basal 1/3 of its length; aedeagal pedon relatively short and wide, and abruptly constricted in apical 1/5, thereafter tapered to rounded point . . Elytra with very minute setae, only perceptible at high magnification; lateral faces of elytra with intervals not noticeably raised; ramus of spermatheca elongate, cylindrical, and slightly thinner than corpus, cornu strongly recurved in basal half with uniquely sinuate appearance, both corpus and cornu with hood-like projections shorter than ramus; males not known . . nodes with bootstrap support above 0.95 are marked with a "*" symbol to the right of each node. In a 697 complementary graph, we show the herein used clade concept labels (Fig. 31).

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The characters, states, and preferred optimizations are described in this section. Characters relating

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The first, classification-based alignment (Fig. 32)    Manuscript to be reviewed Manuscript to be reviewed Figure 31. Preferred phylogeny -clade concept labels. Topology and species-level taxonomic concept labels as in Fig. 30. Clade concept labels, numbered 1-20, are consistently generated by using the alphabetically first epithet in each of the bifurcating sister clades. This method safeguards the clade concept labels against changes due simply to reorientation of leaves. Bold-font square brackets indicate new [JF2018] labels. See also RCC-5 Alignments.