Haemaphysalis vespertina Beati, Egizi & Nava, 2025

Haemaphysalis vespertina Beati, Egizi & Nava , new species ( Figs. 3–6 View FIGURE 3 View FIGURE 4 View FIGURES 5–6 )

ZooBank registration: Details of the new species have been submitted to ZooBank (http://zoobank.org/). The Life Science Identifier (LSID) for the new name Haemaphysalis vespertina is B61B3440-02C5-4944-9221-7B2E703EA237.

Etymology: The specific epithet is derived from the Latin ‘vesper’ in reference to the evening or the evening star and, by extension, to the West. This species is described from the western coast of North America.

Type-locality: USA: California, San Mateo County, Costanoa (coordinates: 37.154342, -122.341978). Collected from vegetation. Known hosts for all stages are Lepus californicus Gray, 1837 and Sylvilagus sp. Gray, 1867; avian hosts such as Melanerpes sp. Swainson, 1832 and Toxostoma sp. Wagler, 1831 have been found infested with immatures.

Female— Figs. 3A–3I View FIGURE 3

HOLOTYPE: USNMENT1785003 ( USA, California, San Mateo County, Costanoa , 37.154342, -122.341978, 9 VII 2024; from vegetation, coll. Tara Roth and Arielle Crews) GoogleMaps . PARATYPES: USNMENT1510975, 5 females ( USA, California, San Mateo County, Costanoa, 37.154342, -122.341978, 9 VII 2024; from vegetation, coll. Tara Roth and Arielle Crews); USNEMNT1510966, 1 female ( USA, California, San Mateo County , Costanoa, 37.154342, -122.341978, 15 VI 2021; from vegetation, coll. Angie Nakano) GoogleMaps . Other material examined: USNEMNT1510970, 22 females ( USA, California, Mendocino County, Hopland, 38.9729541, -123,1163918, 15 VI 1965, ex. L. californicus ); USNMENT1510977, 7 females ( USA, California, Mendocino County , Hopland , 38.9729541, -123.1163918); USNMENT1510986, 1 female ( USA, California, San Benito County , 22 III 1932, ex. L. californicus ); USNMENT1510974, 6 females ( USA, California); USNMENT1510976, 1 female ( USA, Oregon, Harney County , Burns , 43.588333, -119.061389, ex. Sylvilagus sp. ) GoogleMaps .

Body ( Fig. 3A View FIGURE 3 ) of unfed specimens dorsally suboval, longer (1.30–1.49; 1.36 ± 0.06) than wide (0.86–0.95; 0.93 ± 0.03); with lateral edges slightly concave at level of coxa II, widest posterior to mid-length. Scutum oval, ( Fig. 3A View FIGURE 3 , Fig. 3G View FIGURE 3 ) longer (0.79–0.84; 0.82 ± 0.02) than wide (0.62–0.68; 0.64 ± 0.02), with posterior margin rounded; cervical grooves very deep and broad, converging posteriorly to almost mid-length of scutum, broadening posteriorly into shagreened triangular shallower area ( Fig. 3G View FIGURE 3 ); scapulae round, with scattered fine punctations, bearing short, fine setae (0.017–0.027; 0.024 ± 0.003), lateral fields and posterior border with few punctations and glabrous; median field with homogeneously distributed, dense, larger, shallow punctations, all bearing fine setae (0.014–0.025; 0.018 ± 0.003), central punctations sometimes confluent producing rugose effect ( Fig. 3G View FIGURE 3 ). Alloscutum ( Fig. 3A View FIGURE 3 ) with deep, uniformly distributed, very small punctations, all bearing short setae, slightly shorter (0.011–0.022; 0.015 ± 0.003) than scutal setae; marginal groove complete, lining 11 festoons, reaching scutum at level of coxa II; festoons and marginal fold with numerous deep, small, punctations bearing setae. Venter: genital aperture at level of coxae II–III ( Fig. 3E View FIGURE 3 , Fig. 3H View FIGURE 3 ), U-shaped, with almost parallel lateral margins, lined by very narrow sclerotized flaps; anal groove posterior to anus joining genital groove anterolaterally ( Fig. 3C View FIGURE 3 ); in unfed specimens, bean-shaped areas posterolateral to anus, delimited by posteromedian groove, festoons, posterior part of genital groove, and anal groove; ventral grooves more distinct in unfed specimens; punctations dense, fine, deep, uniformly distributed, bearing fine setae (0.011–0.022; 0.016 ± 0.003); spiracular plates almost round with inconspicuous, blunt, dorsal projection, with 2–4 lines of goblet cells, larger in center, slightly smaller along periphery ( Fig. 3F View FIGURE 3 ). Capitulum ( Figs. 3B View FIGURE 3 , Fig. 3D View FIGURE 3 ). Dorsal ( Fig. 3B View FIGURE 3 ): length from palpal apices to tip of cornuae (0.35–0.41; 0.39 ± 0.02); basis capituli broader (0.31–0.35; 0.34 ± 0.01) than long (0.18–0.19; 0.19 ± 0.004), subrectangular, with convex, rounded, lateral edges, posterior margin straight, cornuae wider at insertion than long, rounded; porose areas as narrow flattened ovals (0.050–0.063; 0.055 ± 0.004 long and 0.020–0.035; 0.029 ± 0.04 wide), placed in deep depressions of basis capituli, diverging posteriorly; ventrally ( Fig. 3D View FIGURE 3 ) basis capituli subrectangular, with lateral edges slightly diverging anteriorly, with short, triangular, rounded, posteriorly directed processes, wider at insertion than long. Palps dorsal ( Fig. 3B View FIGURE 3 ): palpal segment I inconspicuous; palpal segment II length (0.16–0.17; 0.16 ± 0.02), palpal segment II width at level of lateral projection (0.13–0.15; 0.14 ± 0.01), distance between apices of lateral projections (0.48–0.54; 0.50 ± 0.02), internal edge of palpal segment II almost straight, with 6 flattened, barbed setae; palpal segment III approximately as long (0.09–0.10; 0.10 ± 0.01) as wide (0.10–0.10; 0.10 ± 0.01); lateral length of palpal segments II and III measured from apex of palpal segment III to tip of angle with lateral projection (0.24–0.26; 0.25 ± 0.001). Palps ventral ( Fig. 3D View FIGURE 3 ): palpal segment I inconspicuous, palpal segment II with no spurs, with approx. 11–12 lanceolate, barbed, flattened median setae, palpal segment III with rounded ventral spur; hypostome clavate, with homogeneous 3:3 dental formula except at crown, approx. 6–7 denticles per file. Legs. Coxa I ( Fig. 3E View FIGURE 3 ) with short, rounded internal spur, wider than long, external spur as triangular ridge, shorter than internal spur, concealed by tuft of long fine, setae; coxa II, III, and IV with single, rounded, triangular spurs, as long as wide at insertion, directed posterolaterally and inserted at mid-width in coxa II and III, directed posteriorly and inserted more medially in coxa IV. Trochanter I with ventral rounded spur; coxae and legs with numerous, very long, fine setae. Haller’s organ as in Fig. 3I View FIGURE 3 .

Male— Figs. 4A–4H View FIGURE 4

ALLOTYPE: USNMENT1785004 ( USA, California, San Mateo County, Costanoa , 37.154342, -122.341978, 9 VII 2024; from vegetation, coll. Tara Roth and Arielle Crews) GoogleMaps . PARATYPES: USNMENT1510975, 4 males ( USA, California, San Mateo County, Costanoa, 37.154342, -122.341978, 9 VII 2024; from vegetation, coll. Tara Roth and Arielle Crews); USNEMNT1510966, 2 males ( USA, California, San Mateo County , Costanoa, 37.154342, - 122.341978, 15 VI 2021; from vegetation, coll. Angie Nakano) GoogleMaps . Other material examined: USNEMNT1510970, 12 males females ( USA, California, Mendocino County, Hopland, 38.9729541, -123,1163918, 15 VI 1965, ex. L. californicus ); USNMENT1510977, 12 males ( USA, California, Mendocino County , Hopland , 38.9729541, -123,1163918); USNMENT1510986, 2 males ( USA, California, San Benito County , 22 III 1932, ex. L. californicus ); USNMENT1785016; 23 males ( USA, California, Bernardino County , 5 IX 1936, ex. L. californicus ); USNMENT1510976, 4 males ( USA, Oregon, Harney County , Burns , 43.588333, -119.061389, ex. Sylvilagus sp. ) GoogleMaps .

Body: conscutum ( Fig. 4A View FIGURE 4 ) distinctly oval, longer (1.16–1.30; 1.21 ± 0.04) than wide (0.72–0.90; 0.80 ± 0.04); with lateral edges slightly convex, widest posterior to mid-length; cervical grooves very deep, short, almost parallel, reaching level of coxa II; scapulae round with scattered fine punctations, bearing short, fine setae as in female; marginal grooves starting anteriorly at approx. mid length of scutum, deep, reaching and lining first festoon, fragmented along other 9 festoons; festoons with scattered punctations and inconspicuous short, fine setae; median field with homogeneously distributed, dense, large, shallow punctations, all bearing short fine setae (0.012–0.019; 0.015 ± 0.002), central punctations sometimes confluent producing distinct rugose effect; lateral fold anterior to festoons with single, lateral, almost linear line of punctations reaching level of coxa II. Venter ( Figs. 4D–F View FIGURE 4 ): genital aperture at level of coxa II, covered by oval apron as in Fig. 4E View FIGURE 4 ; anal groove posterior to anus joining anterolaterally genital groove, bean-shaped areas, posterolateral to anus, delimited by posteromedian groove, festoons, posterior part of genital groove, and anal groove ( Fig. 4D View FIGURE 4 ); punctations dense, fine, deep, uniformly distributed, bearing fine setae (0.014–0.028; 0.020 ± 0.007); spiracular plates almost round with blunt, dorsal projection, with 2–4 lines of goblet cells, larger in center, slightly smaller along periphery ( Fig. 4H View FIGURE 4 ). Capitulum ( Figs. 4B–C View FIGURE 4 ). Dorsal ( Fig. 4B View FIGURE 4 ): length from palpal apices to tip of cornuae (0.25–0.29; 0.27 ± 0.01); basis capituli broader (0.19–0.22; 0.21 ± 0.01) than long (0.10–0.12; 0.11 ± 0.01), subrectangular, with convex, rounded, lateral edges, posterior margin straight, cornuae triangular, wider at insertion than long, rounded; ventrally basis capituli subrectangular, with lateral edges slightly diverging anteriorly, with short rounded, posteriorly directed processes, twice as wide at insertion as long ( Fig. 4C View FIGURE 4 ). Palps dorsal ( Fig. 4B View FIGURE 4 ): palpal segment I inconspicuous; palpal segment II length (0.09–0.10; 0.10 ± 0.001), width at level of lateral projection (0.11–0.12; 0.12 ± 0.001), distance between apices of lateral projections (0.33–0.35; 0.34 ± 0.01); lateral length of palpal segments II and III measured from apex of palpal segment III to tip of angle with lateral projection (0.15–0.16; 0.15 ± 0.004); internal margin concave ending anteriorly with inconspicuous medially directed lobe, with 4 flattened, barbed setae; palpal segment III approximately as long (0.07–0.08; 0.07 ± 0.002) as wide (0.08–0.09; 0.08 ± 0.002). Palps ventral ( Fig. 4C View FIGURE 4 ): palpal segment I inconspicuous, palpal segment II with no spurs, with approx. 9 lanceolate, barbed, flattened median setae; palpal segment III with rounded ventral spur. Hypostome clavate, with homogeneous 3:3 dental formula excepted crown, approx. 6–7 denticles per file. Legs. Coxa I with short, rounded internal spur, wider than long, external spur as rounded ridge, shorter than internal spur, concealed by tuft of long, fine setae; coxa II, III, and IV with single, triangular spurs, approx. as long as wide at insertion level, directed posterolaterally ( Fig. 4F View FIGURE 4 ). Trochanter I with ventral rounded ridge-like spur; coxae and legs with very numerous, long, and fine setae ( Fig. 4F View FIGURE 4 ). Haller’s organ as in Fig. 4G View FIGURE 4 .

Nymph— Figs. 5A–5I View FIGURES 5–6

PARATYPES: USNMENT1510966, 13 nymphs ( USA, California, San Mateo County, Costanoa, 37.154342, -122.341978, 15 VI 2021; from vegetation, coll. Angie Nakano); USNMENT1510980, 4 nymphs ( USA, California, San Mateo County, Costanoa, 37.154342, -122.341978, 15 VI 2021; from vegetation, coll. Angie Nakano). Other material examined: USNMENT1510970, 3 nymphs ( USA, California, Mendocino County, Hopland, 38.9729541, -123,1163918, 15 VI 1965, ex. L. californicus ); USNMENT1785016; 5 nymphs ( USA, California, Bernardino County, 5 IX 1936, ex. L. californicus ); USNMENT1510976, 5 nymphs ( USA, Oregon, Harney County, Burns, 43.588333, -119.061389, ex. Sylvilagus sp. ).

Body: Outline overall oval, length from palpal apices to posterior margin (0.08–0.09; 0.08 ± 0.02); width (0.06– 0.07; 0.07 ± 0.01), with lateral edges slightly concave at level of coxa III, widest at level of coxa IV. Scutum ( Figs. 5C–D View FIGURES 5–6 ) oval, longer (0.45–0.47; 0.46 ± 0.01) than wide (0.43–0.46; 0.45 ± 0.01), with posterior margin rounded; cervical grooves very deep, broad, almost parallel anteriorly, broadening posteriorly into triangular shallower area and reaching edge of scutum; scattered, unevenly distributed punctations bearing fine, short setae (0.007–0.020; 0.013 ± 0.003). Alloscutum ( Fig. 5E View FIGURES 5–6 ) with posteromedian and curved, posterolateral grooves outlined by very fine, deep, glabrous, dense punctations; other areas of idiosoma with sparse, scattered, medium-sized punctations bearing setae (0.009–0.019; 0.013 ± 0.002); 11 festoons, with medially directed 1 or 2 setae each (not on central festoon); marginal groove lining all festoons and reaching scutum at level of coxae III. Venter ( Figs. 5F–G View FIGURES 5–6 ): anal groove posterior to anus, median postanal groove reaching festoons, joining anterolaterally hint of future genital grooves that extend posteriorly to festoon level ( Fig. 5F View FIGURES 5–6 ); scattered, sparse, small punctations bearing fine setae (0.020–0.038; 0.026 ± 0.004); spiracular plates sub-oval with 1–3 lines of goblet cells, larger in center, smaller in peripheral line ( Fig. 5H View FIGURES 5–6 ). Capitulum ( Figs. 5A–5B View FIGURES 5–6 ). Dorsal: length from palpal apices to tip of cornuae 0.25; 0.24 ± 0.01); basis capituli broader (0.18–0.18; 0.18 ± 0.001) than long (0.09–0.1; 0.09 ± 0.002), subrectangular, with convex, rounded, lateral edges, posterior margin convex, cornuae wider at insertion than long, triangular, and pointed ( Fig. 5B View FIGURES 5–6 ); ventral basis capituli subrectangular with lateral edges diverging anteriorly, with distinct, rounded, posterolaterally directed processes ( Fig. 5A View FIGURES 5–6 ). Palps dorsal ( Fig. 5B View FIGURES 5–6 ): palpal segment I inconspicuous; palpal segment II length (0.08–0.09; 0.08 ± 0.003), palpal segment II width at level of lateral projection (0.09–0.10; 0.10 ± 0.003), distance between apices of lateral projections (0.28–0.30; 0.29 ± 0.006), internal edge of palpal segment II almost straight, with 2 flattened, barbed setae; palpal segment III approximately as long (0.06–0.07; 0.07 ± 0.003) as wide (0.06– 0.07; 0.06 ± 0.003); lateral length of palpal segments II and III measured from apex of palpal segment III to tip of angle with lateral projection (0.12–0.13; 0.13 ± 0.003). Palps ventral ( Fig. 5A View FIGURES 5–6 ): palpal segment I inconspicuous, palpal segment II with no spurs, with 5 lanceolate, barbed, flattened median setae, palpal segment III with rounded ventral spur. Hypostome clavate, with homogeneous 2:2 dental formula excepted crown, approx. 7 denticles per file. Legs. Coxa I ( Fig. 5G View FIGURES 5–6 ) with rounded internal spur, as wide as long, external spur as inconspicuous ridge, concealed by long, fine seta; coxa II, III, and IV with single, rounded, triangular spurs, directed posterolaterally and inserted at mid-width in coxa II and III, directed posteriorly in coxa IV. Trochanter I with ventral triangular spur; coxae and legs with numerous, 2–3 very long, fine setae. Haller’s organ as in Fig. 5I View FIGURES 5–6 .

Larva— Figs. 6A–6G View FIGURES 5–6

PARATYPES: USNMENT1510966, 2 larvae ( USA, California, San Mateo County, Costanoa , 37.154342, - 122.341978, 15 VI 2021; from vegetation, coll. Angie Nakano) GoogleMaps ; USNMENT1510991, 1 larva ( USA, California, San Mateo County, Costanoa , 37.154342, -122.341978, 15 VI 2021; from vegetation, coll. Angie Nakano) GoogleMaps . Other material examined: USNMENT1510964, 1 larva ( USA, California, Los Angeles County, 13 XI 1987, ex. Toxostoma redivivum , sent by K.C. Emerson); USNMENT1510958, 5 larvae ( USA, California, San Diego County, Imperial Beach, 21 XII 1968, ex. Toxostoma bendirei ); USNMENT1510962, 8 larvae ( USA, California, Sonoma County , Jack London State Historic Park, 38.350556, -122.543056, 24 IX 2021, ex. vegetation, coll. Megan Saunders); USNMENT 1510970, 50 larvae ( USA, California, Mendocino County , Hopland, 38.9729541, -123,1163918, 15 VI 1965, ex. Lepus californicus ); USNMENT1785017, 1 larva ( USA, California, Yalo County , Winters, 38.525, - 121.970833, XII 1965, ex. Melanerpes formicivorus ) GoogleMaps .

Body: ( Fig. 6C–D View FIGURES 5–6 ): dorsally subcircular, lateral margins slightly concave at level of leg 2, length from tip of scapulae to posterior edge (0.45–0.50; 0.47 ± 0.02), widest (0.40–0.46; 0.43 ± 0.02) near midlength, median area of idiosoma posterior to scutum convex; 11 festoons. Scutum (Fig: 6C): length (0.24–0.25; 0.25 ± 0.002), breadth (0.30–0.32; (0.31 ± 0.06), outline broadly cordiform; scapulae rounded; cervical grooves almost parallel, distinct, deep, wide, with margins slightly diverging posteriorly, reaching 1/3 of scutal length; cervical fields wide, very shallow, reaching edge of scutum, delimiting slightly convex posterior central field and convex anterolateral scutal areas; 3 pairs of scutal setae (0.009–0.014; 0.012 ± 0.002) and 4 pairs of small wax glands. Alloscutum ( Fig. 6D View FIGURES 5–6 ): large wax glands (or sensilla sagittiformia) present; 8 pairs of fine dorsomarginal setae (0.014–0.022; 0.017 ± 0.002), two anterior to large wax glands; 2 pairs of central dorsal setae (0.009–0.014; 0.011 ± 0.002); supplementary setae absent. Venter ( Fig. 6E–F View FIGURES 5–6 ) with 3 pairs of large wax glands posterior to each coxa; 3 pairs of sternal setae (0.016–0.024; 0.020 ± 0.002), two aligned with coxae III and one with coxae II; 2 pairs of preanal setae (0.010–0.020; 0.014 ± 0.004); 4 pairs of premarginal setae (0.011–0.016; 0.013 ± 0.002), 4 pairs of marginal ventral (0.013–0.018; 0.015 ± 0.002), and 1 pair of minuscule anal setae. Capitulum ( Fig. 6A–B View FIGURES 5–6 ): dorsal length ( Fig. 6A View FIGURES 5–6 ) from palpal apices to tip of cornua (0.13–0.15; 0.14 ± 0.07), width between tips of lateral extensions of palpal segments II (0.15–1.60; 0.16 ± 0.03). Basis capituli length from papal insertion to tip of cornua (0.06–0.07; 0.07 ± 0.002), width (0.11–0.12; 0.12 ± 0.01), with dorsal posterior edge concave, lateral margins rounded, with small notch under insertion of palps, cornua short, wider at insertion than long, bluntly pointed and posteriorly directed. Palpal segment I inconspicuous, palpal segment II almost as long (0.05–0.05; 0.05 ± 0.006) long as broad (0.05– 0.05; 0.05 ± 0.006) (width measured at level of lateral extensions), palpal segment III (0.03–0.03; 0.03 ± 0.001) long by (0.04–0.04; 0.04 ± 0.001) broad. Palpal segments II and III not fused, delimited by distinct dorsal suture; palpal segment II dorsally with deep groove going from posteromedian edge to tip of lateral protrusion and with 1 medially directed flattened barbed seta. Ventrally ( Fig. 6B View FIGURES 5–6 ), basis capituli broadly rectangular, with posterolaterally directed, bluntly pointed, triangular auriculae. Hypostome spatulate, toothed portion covering approx. 3⁄4 of hypostomal length; dental formula below crown 2:2 on 4–5 rows, denticles of approx. similar size; one pair of post-hypostomal short and fine setae. Palpal segment II with 2 long, flattened, barbed setae inserted on median edge and covering in part hypostome and post-hypostomal setae; palpal segment III with rounded subtriangular ventral spur; palpal segment IV extruding, with approx. 7 apical setae. Palpal setation as in Fig. 6A–B View FIGURES 5–6 . Legs: Coxa I with internal broad, rounded spur; coxa II with one, short, wide, rounded spur; coxa III with a ridge-like spur; coxal setae approx. twice as long as idiosomal setae (0.03–0.05; 0.04 ± 0.005) ( Fig. 6E View FIGURES 5–6 ). Haller’s organ as in Fig. 6G View FIGURES 5–6 .