Whetwhetaksa millerae Archibald & Cannings, 2021

Archibald, Bruce, Cannings, Robert A., Erickson, Robert J., Bybee, Seth M. & Mathewes, Rolf W., 2021, The Cephalozygoptera, a new, extinct suborder of Odonata with new taxa from the early Eocene Okanagan Highlands, western North America, Zootaxa 4934 (1), pp. 1-133 : 111-112

Whetwhetaksa millerae Archibald & Cannings , new species

Fig. 66–70 .

Diagnosis. As for genus and family.

Type and additional material. All are isolated wings. Holotype: SR 06-01-38 ( Fig. 66 ): Republic B4131 collected by Gregg Wilson, 13.vi.2005; paratype 1: SR 10-36-10 ( Fig. 67 ), Republic B4131, collected by Shannon Miller, June. vi.2010; paratype 2: SRUI 99-98-34 ( Fig. 68 ), Republic B4131 collected by Shannon Miller, 6.x.2011. Additional material: SR 93-11-01 ( Fig. 69 ), Republic A0307 B collected by unknown person, 15.vii.1986 ; SR 06-01-36 ( Fig. 69 ), collected at Republic B4131 by Gregg Wilson, 18.vi.2004; SR 05-11-01AB ( Fig. 70 ), Republic A0307 collected by Peg Johnson, 11.vi.2005 ; SR 10-14-19 ( Fig. 70 ), Republic B4131 collected by Stonerose staff, 23.iv.2010, a wing fragment of the anterior mid-wing. All specimens are housed in the collections of the Stonerose Interpretive Center .

Description. Holotype, SR 06-01-38, wing ( Fig. 66 ). Mostly complete with small sections obscured, missing. Length 31.8 mm, arculus to apex: 27.9 mm, nodus to apex: 20.4 mm, origin of RP2 to apex: 17.5 mm, arculus to base of pterostigma: estimated 19.5 mm, nodus to base of pterostigma: estimated 11.8 mm, nodus at 44% wing length, width 8 mm. Infuscate in proximal half (in well-preserved portions), apical half hyaline (sex unknown). Pterostigma dark, estimated 5–5.5 mm long, eight–nine times longer than wide (distal end damaged), no oblique brace vein, subtending seven cells. Preserved crossveins in postnodal, postsubnodal spaces not aligned: 13 postnodal (or portions) preserved (likely five to six? more total); 17 postsubnodal preserved to pterostigma, likely few more total. Two cells wide distal to pterostigma in costal space. RA meets margin at apex. IR1 origin not preserved; RP2 origin 3.5 cells of RP1-2–IR2 space distal to subnodus; IR2 origin about 3/5 arculus to nodus; IR2 close to linear, only slightly zigzagged in preserved portions. RP3-4 origin>2/5 distance arculus to nodus. Arculus close to, just distal to Ax1. No accessory antenodal crossveins. Ax0 not detected. No crossvein O. Quadrangle about three times longer than wide, distal side twice length of proximal, anterior side about 5/6 length of posterior. Nodal, subnodal crossveins with normal obliquity. MA zigzags distal to about mid-wing (region poorly preserved); MP apparently linear from quadrangle to terminus (but distal portion poorly preserved), gently curved. CuA long, ends on margin just distal to mid-wing (about mid-way arculus to apex), well distal to base of RP2. CuA slightly zigzagged distal to level of quadrangle, more so at terminus (but much of distal half poorly or not preserved); CuA–A space partly missing, two cells wide preserved, at least three, perhaps four total.

Paratype 1, SR 10-36-10, wing ( Fig. 67 ). Mostly whole, missing the proximal posterior region and a region of the posterior-mid region in the proximal part of the apical third. Length 30.5 mm (proximal-most part missing); arculus to apex: 26.8 mm; nodus to apex: 18.8 mm; origin of RP2 to apex: 16.5 mm; arculus to base of pterostigma: 18.1 mm; nodus to base of pterostigma: 10.1 mm; nodus at 40% wing length; width 8.0 mm. Colouration well-preserved (sex unknown): all preserved portions of proximal half (base to postnodal crossvein 6) darkly infuscate, apical half very lightly infuscate. Pterostigma dark, distal end oblique, 5.3 mm long, length about 11.5 times width, subtends 7.5 cells. No oblique brace vein; 13 crossveins preserved in postnodal space (probably one more), 17 in postsubnodal space to pterostigma (total), only one-two aligned. Two cells wide in costal space distal to pterostigma. RP1 meets margin at apex. RP1 space becomes two cells wide in level of pterostigma (region poorly preserved); IR1 origin slightly zigzagged, six cells distal to origin of RP2 ; IR1 two cells wide distal to level of base of pterostigma (region poorly preserved); RP2 origin 2.5 RP1-2 –IR2 space cells distal to subnodus. IR2 origin about 3/5 distance arculus to subnodus; RP3-4 origin>2/5 distance arculus to nodus. Arculus close to, distal to Ax1. Ax0 not detected. No accessory antenodal crossvein detected (but region where one might be not preserved). No crossvein O. Quadrangle region not preserved. Nodal crossvein, subnodus angled as in holotype. MA linear in all preserved portions (some distal to mid-wing not preserved); MP preserved distal to between levels of origins of RP3-4 , IR2: linear to terminus. CuA linear, deeply curved, wide; CuA–A space six cells wide not at widest (maybe one–two more total?) .

Paratype 2, SR 99-98-34, wing ( Fig. 68 ). Missing the posterior portion except in the apical quarter. Length 35.8 mm (proximal-most part missing), nodus to apex: 20.7 mm, origin of RP2 to apex: 19.5 mm, nodus to base of pterostigma: 12.4 mm; nodus at 43% wing length; width unknown by preservation. Colouration well-preserved (sex unknown): all preserved regions of proximal half (base to postnodal crossvein 6) darkly infuscate, apical half lightly infuscate. Pterostigma anterior, posterior margins oblique, posterior more so, length 5.4 mm, length about 9.5 times width, subtends eight cells. No oblique brace vein; nodal crossvein, subnodus oblique as in holotype, 18 crossveins in postnodal space (total), 19 postsubnodal space (probably one more), one-two aligned. Two cells wide in most of costal space distal to pterostigma. RA meets margin at apex. RP1 –IR1 space becomes two cells wide at level of mid-pterostigma. IR1 origin not preserved, slightly zigzagged until near margin, then breaks up in strong zigzag. IR1– RP2 space becomes two cells wide same level as RP1 –IR1 space. RP2 origin close to subnodus, about one cell distant. IR2 origin not preserved, but far proximal to subnodus (could be as in holotype); origin, most of RP3-4 not preserved); arculus, quadrangle not preserved; Ax0 not detected; one accessory antenodal crossvein; region where crossvein O might be not preserved; nothing proximal to fragments of RP3-4 preserved .

Additional material.

SR 93-11-01, wing ( Fig. 69 ). Anterior mid-portion. Length: nodus to base of pterostigma: 11.3 mm. Colouration difficult to assess by preservation, possibly like holotype (sex unknown); 16 crossveins preserved in postnodal space (probably one more total), 17 in postsubnodal space (total), only one–two aligned. IR1 origin seven cells RP1 – RP2 space distal to origin of RP2 ; RP2 origin 2.5 cells RP1-2 –IR2 space distal to subnodus; arculus not preserved, but IR2, RP3-4 origins appear the same position proximal to subnodus as in holotype, paratypes 1, 2; no crossvein O (in region preserved); MA, MP, CuA linear in all portions preserved; little of CuA–A space preserved, at least three cells wide .

SR 06-01-36, wing ( Fig. 69 ). Much of the basal half (unknown if fore- or hind) missing proximal-most portion. Colouration difficult to determine by preservation (sex unknown). Few crossveins preserved in postnodal, postsubnodal spaces, none aligned. Subcosta may cross nodus (but could be adventitious crossvein between costa, nodus?). IR2, RP3-4 origins as in holotype, paratypes, roughly mid-way between arculus, subnodus. Nodal, subnodal crossveins of normal obliquity. Arculus far-proximal to Ax2 as in holotype, paratypes (Ax1 not preserved). Quadrangle similar to holotype. All main vein portions preserved (proximal portions of RP1-2 , IR2, RP3-4 , MA, MP, CuA) linear. CuA–A space with four cells wide preserved (one or more further complete?) .

SR 05-11-01AB, wing ( Fig. 70 ). Mostly complete, very crushed, torn, apical region bent. Length: arculus to base of pterostigma: 23? mm, nodus to base of pterostigma: 11.3 mm. Colouration (sex unknown): proximal half apparently darkly infuscate throughout as preserved, apical half apparently very lightly infuscate as preserved. Pterostigma little-preserved; 20 crossveins preserved in postnodal space (likely one–two? more), 15 in postsubnodal space, likely many more?, one–two aligned. Arculus closer to, just distal to Ax1. Ax0 detected at wing base. One accessory antenodal crossvein detected. Crossvein O not detected. Major veins very broken up, crimpled; all preserved portions linear. CuA–A space with four cells at widest (level of nodus), perhaps similar to SR 06-01-36.

SR 10-14-19, wing ( Fig. 70 ). Small anterior mid-portion. Proximal portion darkly infuscate as in paratypes 1, 2, apical portion hyaline as preserved (sex unknown). Nodus, subnodus angled as in other specimens. Origins of RP2 , IR2, crossvenation in postnodal, subnodal spaces as in holotype .

Etymology. The specific epithet is a patronymic formed from the surname of Shannon Miller, who found paratypes 1 and 2.

Range and age. Republic localities B4131 and A0307B of the Tom Thumb Tuff Member of the Klondike Mountain Formation; latest Ypresian.

Discussion. There is notable variation among these specimens in the length of the antenodal space, the presence of an accessory antenodal crossvein, and the width of the CuA–A space. SR 99-98-34 has a longer antenodal space measured Ax1 to nodus of 11 mm. In SR 10-36-10 this is 8.8 mm; in SR 06-01-38, 8.0 mm; and in SR 05-11-01, 9.0 mm. SR 05-11-01, SR 99-98-34 each possess a single accessory antenodal crossvein. In SR 10-36-10 and SR 10-14-19, this is not determinable by preservation, whereas SR 06-01-38, SR 06-01-36, and SR 93-11-01 definitely do not have this crossvein. SR 10-36-10 has a much wider CuA–A space (at least six cells deep; probably more: the widest part is not preserved), whereas SR 06-01-38, SR 06-01-36 and SR 05-11-01 apparently have a maximum of four cells deep (others not known by preservation). Forewing/hind wing differences and sexual dimorphism might explain some or all of this variation; more complete specimens are needed to evaluate these possibilities and so at present we refrain from assuming more than one species here.

Figures

FIGURE 66. Whetwhetaksa millerae holotype, SR 06-01-38. A, photograph; B, drawing. Both to scale, 5 mm.

FIGURE 67. Whetwhetaksa millerae paratype 1, SR 10-36-10. A, photograph; B, drawing. Both to scale.

FIGURE 68. Whetwhetaksa millerae paratype 2, SRUI 99-98-34. A, photograph; B, drawing. Both to scale; aa, accessory antenodal crossvein.

FIGURE 69. Whetwhetaksa millerae additional material. SR 93-11-01: A, photograph; B, drawing; SR 06-01-36: C, photograph; D, drawing.All to scale, 5 mm.

FIGURE 70. Whetwhetaksa millerae, additional material. SR 10-14-19: A, photograph; B, drawing; SR 05-11-01: C, photograph; D, drawing.All to scale, 5 mm.