Waynokiops Hill, Pfeiffer & Jacobus 2010

Kluge, Nikita, 2022, Two new species of Waynokiops Hill et al. 2010 (Ephemeroptera: Baetidae) from the Oriental Region, Zootaxa 5182 (1), pp. 41-63 : 42-44

Waynokiops Hill, Pfeiffer & Jacobus 2010

( Figs 1–63 , 65–94 )

Type species: Waynokiops dentatogriphus Hill, Pfeiffer & Jacobus 2010 .

Diagnosis and comparison

Larva:

(1) Head is narrowed, with frontal suture acute-angled ( Fig. 15 ; Hill et al., 2010: fig. 1).

(2) Labrum is trapezoidal, i.e. widened distally, with lateral margins diverging and distal margin more straight than in other Baetidae ; median emargination is widened and shallow ( Figs 26–27 , 65–67 ; Hill et al. 2010: fig. 2). At least in W. pentacanthus sp. n. and W. palifer sp. n., bottom of median emargination is straight; 4 sensilla form transverse row on ventral side of labrum proximad of the median emargination ( Figs 27 , 66 ) (in contrast with the majority of Baetidae , where the bottom of the median emargination forms a sclerotized projection). Stout setae, forming regular row on distal margin, are spaced (in contrast to most other Baetidae , in which sockets of these setae are contiguous). At least in W. pentacanthus sp. n. and W. palifer sp. n., dorsal surface of labrum bears only irregularly situated fine setae, without distinguishable submedian pair and without distinguishable pair of submarginal rows. Shape and setation of labrum are similar to that of Securiops ( Jacobus et al. 2006: fig. 7).

(3) Mandibles are of the « Centroptilum - type », i.e. with the kinetodontium separated from the incisor (but remains to be immovable) and rotated perpendicular to the plane of the mandible ( Figs 21–22 , 33–34 ; Hill et al. 2010: figs 3–4). The same in some other taxa ( Kluge 2020b: 440).

(4) Maxilla is of the « Cloeon - type », i.e. with all three canines and three dentisetae long, slender and bent in the same direction. Outer side laterad of canines lacks setae ( Figs 23 , 68 ; Hill et al. 2010: fig. 5). The same in other Cloeon /fg1.

(5) Maxillary palp is long and 2-segmented ( Figs 23 , 68 ; Hill et al., 2010: fig. 5).

(6) Labium has glossae and paraglossae equally wide ( Fig. 24 ; Hill et al. 2010: fig. 6), with setation of the « Cloeon - type » ( Fig. 70 ).

(7) Distal (3rd) segment of labial palp is widened distally and truncate (as in other Cloeon /fg1).

(8) Distal (3rd) segment of labial palp is especially wide, with the inner-distal angle acute; the outer-distal angle is sharply stretched into a small, blunt projection ( Figs 24 , 71 ; Hill et al. 2010: fig. 7). This resembles Securiops Jacobus et al. 2006 , which has pointed projection on the same place.

(9) Femora have no any stout setae on outer side and apex (in contrast to other Baetidae ); inner side of femur with row of stout setae ( Figs 37 , 79 ).

(10) Tibiae also have no any stout setae on outer side and apex, with row of stout setae on inner side ( Figs 35 , 39–40 , 79 ). Patella-tibial suture varies among species (see below).

(11) Claw is long and slender, slightly arched, symmetric, with two equal rows of denticles ( Figs 36 , 79 , 87 ; Hill et al., 2010: fig. 11) (this structure is primitive for Baetidae ).

(12) Abdominal terga and other body parts without scales and scale sockets (at least in W. pentacanthus sp. n. and W. palifer sp. n.) ( Figs 3–9 , 88–91 ) (in contrast to majority of other Cloeon /fg1 and majority of Baetidae ).

(13) Each abdominal tergum II–VI bears unpaired dorsal protrusions hooked backward ( Figs 3–4, 10 , 72 , 88–89 ; Hill et al., 2010: fig. 9).

(14) Posterior abdominal segments, at least segments 8–9, bear denticles on lateral margins ( Figs 28 , 72 ) (that is characteristic for Cloeon /fg1).

(15) Tergalii of certain pairs with additional dorsal lamella; the dorsal lamella arises as an expansion of the anal margin of tergalius, is bent dorsally and overlaps the tergalius from dorsal side ( Figs 28–32 , 75–78 , 94 ; Hill et al., 2010: fig. 12) (this is characteristic for Cloeon /fg1). Tergalii are capable of rhythmic respiratory movements (as in other Cloeon /fg1).

(16) All three caudalii (cerci and paracercus) are equally long, with rows of primary swimming setae on inner sides of the cerci and outer sides of the paracercus; there are no secondary swimming setae on lateral sides of cerci ( Figs 11, 19 ) (that is primitive for Baetidae and Ephemeroptera in general).

(17) Posterior margin of each 4th segment of cercus and paracercus darkened and bears enlarged denticles ( Figs 11, 19 ) (that is characteristic for Cloeon /fg1).

Wings:

(18) Hind wings are completely absent; larval metanotum lacks vestiges of hind protoptera.

Male genitalia (known for W. pentacanthus sp. n. only):

(19) Before molt to subimago, subimaginal gonostyli are bent under the larval cuticle in the « Cloeon - type » pose, i.e. with 2nd segments directed laterally ( Fig. 56 ).

(20) Sterno-styligeral muscle is retained ( Fig. 58 ).

(21) Gonovectes are fused with penial bridge ( Fig. 58 ) (as in other Cloeon /fg1).

(22) Imaginal gonostyli are similar to that of other Cloeon /fg1: with 2nd segment widened at apex, 3rd segment small and petiolate ( Figs 53 , 58 ).

Among these characters, the following are apomorphies of Waynokiops :

(2) median emargination of labrum widened, without projection, with transverse row of four sensilla on ventral side (the same in Securiops );

(8) distal segment of labial palp with outer-distal angle stretched into peculiar narrow projection (similar to that of Securiops );

(9) outer margin of the femur without stout setae;

(12) abdominal terga without scales and scale sockets (the same in Securiops );

(13) each abdominal segment II–VI with a medial dorsal hooked protrusion.

Key to larvae of Waynokiops

1(2) Abdominal segment VII with pair of wide, triangular lateral expansions similar to that of previous uromeres ( Hill et al. 2010: fig. 8).................................................................................. W. dentatogriphus

2(1) Abdominal segment VII without lateral expansions ( Figs 28 , 72 )................................................ 3

3(4) Tergalius II without dorsal lamella ( Fig. 74 , 93 ); median protrusion of abdominal tergum II widened distally ( Fig. 72 , 88 )...................................................................................... ..... W. palifer sp. n.

4(3) Tergalius II with dorsal lamella ( Fig. 29 ); median protrusion of abdominal tergum II narrowed distally ( Figs 3 , 28 )........................................................................................... W. pentacanthus sp. n.

Figures

FIGURES 1–10. Waynokiops pentacanthus sp. n., larvae. 1–2, female larvae of last instar; 3–8, median part of abdominal terga II, VI, VII, VIII, IX and X; 9, median part of sternum VII; 10, thorax and abdomen of young larva.

FIGURES 11–20. Waynokiops pentacanthus sp. n., larval exuviae (11–18 with same magnification). 11, abdomen; 12–14, half of pronotum and mesonotum (arrows show small proruberances); 15, head; 16–18, fore, middle and hind legs; 19, fragment of cercus (dorsal view); 20, paraproct (11, 14, 16–19, holotype).

FIGURES 21–27. Waynokiops pentacanthus sp. n., larvae. 21–22, left and right mandibles; 23, maxilla; 24, labium (focus on ventral side); 25, hypopharynx with superlinguae; 26, labrum (focus on dorsal side); 27, the same, middle of anterior margin (focus on ventral side).

FIGURES 28–34. Waynokiops pentacanthus sp. n., larvae. 28, abdomen without tergalii II–V; 29–32, tergalii II–V; 33–34, apices of right and left mandibles.

FIGURES 35–38. Waynokiops pentacanthus sp. n., exuviae of larval fore legs. 35–36, two fragments of tibia, tarsus and claw, ventral view; 37, femur; 38, apex of femur and base of tibia. Abbreviation: pts, patella-tibial suture crossing ventral side of tibia.

FIGURES 39–42. Waynokiops pentacanthus sp. n., patella-tibial suture, anterior view. 39–40, fore and middle legs of larva; 41–42, fore and middle legs of female subimago.

FIGURES 43–52. Waynokiops pentacanthus sp. n. 43–45, male imagines; 46, fore leg of male imago; 47, fore leg of female imago; 48, its subimaginal exuviae; 49, middle leg of female imago; 50, female imago; 51, subimaginal exuviae of meso- and metapleuron; 52, subimaginal exuviae of mesonotum (44–46, holotype).

FIGURES 53–55. Waynokiops pentacanthus sp. n. 53, genitalia of male imago, focus on dorsal side; 54, their subimaginal exuviae; 55, genitalia of male imago, focus on ventral side (holotype).

FIGURES 56–59. Waynokiops pentacanthus sp. n. 56, precursors of subimaginal gonostyli starting to develop under larval cuticle; 57, subimaginal exuviae of gonostylus; 58, genitalia of male imago; 59, wing. Abbreviations: g1–g3, segments of gonostylus; gd, gonoduct; m.IX-X, areas of anterior attachment of paired intersegmental muscles going to paraprocts base; m.s, median sterno-styligeral muscle.

FIGURES 60–63. Waynokiops spp., eggs. 60–61, W. pentacanthus sp. n.; 62–63, W. palifer sp. n.

FIGURES 65–71. Waynokiops palifer sp. n., larva. 65–66, anterior margin of labrum, with focus on dorsal and ventral sides; 67, outline of labrum; 68, maxilla; 69, apex of left mandible; 70, glossae and paraglossae (at left dorsal view, at right ventral view); 71, 2nd and 3rd segments of labial palp.

FIGURES 72–80. Waynokiops palifer sp. n., larva. 72, abdomen without tergalii II–VI; 73, enlarged posterolateral spine of uromere VI; 74–78, tergalii II–VI; 79, hind leg with subimaginal leg inside; 80, paraproct.

FIGURES 81–87. Waynokiops palifer sp. n. 81, head of female subimago in larval cuticle; 82–83, thorax and abdomen of female subimago extracted from larval cuticle, ventral and dorsal view; 84–86, larval fore, middle and hind legs with subimaginal legs inside; 87, proximal part of claw.

FIGURES 88–94. Waynokiops palifer sp. n., larval exuviae. 88–90, fragments of abdominal terga II, VI and IX; 91, fragment of female sternum IX; 92–94, tergalii I, II and IV.