Wayne P Maddison1. 1. Beaty Biodiversity Museum and Departments of Zoology and Botany, University of British Columbia, 6270 University Boulevard, Vancouver, British Columbia, V6T 1Z4, Canada.
Abstract
The harmochirine jumping spiders include the New World Habronattus, notable for their complex courtship displays, and Pellenes, found throughout the Old World and North America. Five new species of Habronattus and one new species of Pellenes are here described from North America: Habronattus aestus, sp. n., Habronattus chamelasp. n., Habronattus empyrussp. n., Habronattus luminosussp. n., Habronattus robertisp. n., and Pellenes canadensissp. n. For each of the new species, photographs of living specimens are given, as well as notes on habitat. The new subgenus Pellenattus is described for the subgroup of Pellenes restricted to North America, with type species Pellenes peninsularis Emerton, 1925. Species placed in Pellenes (Pellenattus) are Pellenes apacheus Lowrie & Gertsch, 1955, Pellenes canadensissp. n., Pellenes crandalli Lowrie & Gertsch, 1955, Pellenes dorsalis (Banks, 1898b), Pellenes grammaticus Chamberlin 1925, Pellenes levii Lowrie & Gertsch, 1955, Pellenes limatus Peckham & Peckham, 1901, Pellenes longimanus Emerton, 1913, Pellenes peninsularis Emerton, 1925, Pellenes shoshonensis Gertsch, 1934, and Pellenes washonus Lowrie & Gertsch, 1955. Pellenes wrighti Lowrie & Gertsch, 1955 is synonymized with Pellenes peninsularis. Attention is drawn to an undescribed species of Habronattus from Canada whose only known specimen is apparently lost.
The harmochirine jumping spiders include the New World Habronattus, notable for their complex courtship displays, and Pellenes, found throughout the Old World and North America. Five new species of Habronattus and one new species of Pellenes are here described from North America: Habronattus aestus, sp. n., Habronattus chamelasp. n., Habronattus empyrussp. n., Habronattus luminosussp. n., Habronattus robertisp. n., and Pellenes canadensissp. n. For each of the new species, photographs of living specimens are given, as well as notes on habitat. The new subgenus Pellenattus is described for the subgroup of Pellenes restricted to North America, with type species Pellenes peninsularis Emerton, 1925. Species placed in Pellenes (Pellenattus) are Pellenes apacheus Lowrie & Gertsch, 1955, Pellenes canadensissp. n., Pellenes crandalli Lowrie & Gertsch, 1955, Pellenes dorsalis (Banks, 1898b), Pellenes grammaticus Chamberlin 1925, Pellenes levii Lowrie & Gertsch, 1955, Pellenes limatus Peckham & Peckham, 1901, Pellenes longimanus Emerton, 1913, Pellenes peninsularis Emerton, 1925, Pellenes shoshonensis Gertsch, 1934, and Pellenes washonus Lowrie & Gertsch, 1955. Pellenes wrighti Lowrie & Gertsch, 1955 is synonymized with Pellenes peninsularis. Attention is drawn to an undescribed species of Habronattus from Canada whose only known specimen is apparently lost.
The two jumping spider genera F.O. Pickard-Cambridge, 1901 and Simon, 1876 are closely related within the subtribe (Maddison and Hedin 2003a, b; Maddison 2015). While species are confined to the Americas (Griswold 1987) and known for their complex courtship ornamentation and behaviour (Peckham and Peckham 1890; Griswold 1987; Elias et al. 2003, 2012), are distributed throughout the Old World, along with North America, and show considerably less sexual dimorphism and courtship complexity. The phylogeny of has been studied by both morphological (Griswold 1987) and molecular data (Maddison and Hedin 2003b), but many ambiguities remain — thus, an ongoing phylogenetic study seeks to use genomic data. To offer names for 6 taxa used in that phylogenomic study, five new species of and one new species of are described from North America. In addition, a new subgenus is erected to house the North American group of .
Methods
Specimens are deposited in the , the , the , or the .Spencer Entomological Museum of the University of British ColumbiaColección Nacional de Arácnidos, Instituto de Biología, Universidad Nacional Autónoma de MéxicoMuseum of Comparative Zoology, Harvard UniversityAmerican Museum of Natural HistoryPreserved specimens were examined under both dissecting microscopes and a compound microscope with reflected light. Drawings (except that of habitus and palpi from Ontario) were made with a drawing tube on a Nikon ME600L compound microscope.Terminology is standard for . The descriptions were written with primary reference to the focal specimen indicated, which was used for measurements and carefully checked for details, but they apply as far as known to the other specimens examined. All measurements are given in millimeters. Carapace length was measured from the base of the anterior median eyes not including the lenses to the rear margin of the carapace medially; abdomen length to the end of the anal tubercle. Rotation of the bulb of the palp expressed in degrees counterclockwise from distal. Thus, 0° is distal (12:00 on an analog clock face); 90° is prolateral (9:00); 180° is proximal (6:00); 270° is retrolateral (3:00). The following abbreviations are used: AME; ALE; PLE; PME; RTA. The apophysis accompanying the embolus of the male palp was called the conductor by Lowrie and Gertsch (1955), the tegular apophysis by Griswold (1987), and the compound terminal apophysis by Logunov et al. (1999). It is here called the “terminal apophysis”, abbreviated “TmA”, following Edwards (2015)., anterior median eyes, anterior lateral eyes, posterior lateral eyes, posterior median eyes (the “small eyes”), retrolateral tibial apophysis
Taxonomy
F. O. Pickard-Cambridge, 1901
Type species.
Peckham & Peckham, 1896
Notes.
Most of the approximately 100 species (Griswold, 1987) of are found in Mexico and the United States, extending into arctic Canada and to southern South America. as a whole is easily recognized by the 90° bend (“elbow”) on the long thin TmA of the palp, though the elbow is lost secondarily in the species group. Several clades of species are recognized as species groups (Griswold 1987; Maddison and Hedin 2003b), some referred to here in the descriptions.Two of the new species were studied by Maddison and Hedin (2003b): , called “ sp. (CHMLA)” by Maddison & Hedin, and , called “ sp. (ROBRT)”. For the other new species we used informal names in field and lab notebooks: as “peñasco” or “ESTU”, as “blondie” or “BLNDI”, as “” or “SUNGL”, and as “ cf. ”. Images and mentions of have appeared on news reports in connection with Zurek et al.’s (2015) study of colour vision, under the name “”.Maddison
sp. n.http://zoobank.org/43D988D1-AA30-45F6-8A90-15D67B59EAA7Figs 1–12
From the Latin, in reference to the tides and the heat of its habitat.
Diagnosis.
This species can be placed in the group by male ornamentation (shelf of projecting setae under the AME, Fig. 7; darkened first tarsus, Fig. 6) and the relatively short and pointed terminal apophysis (TmA) arising at about 120°. However, unlike other members of the group (Griswold 1987, figures 184–186), the TmA of is thin at the base (Fig. 1), more or less lacking the elbow typical of . The male’s scantiness of ornamentation is also distinctive — the first males found were not recognized as adult initially — as is the habitat of saline negative estuaries. Both male and female have the first tibia reddish brown, contrasting against a darker patella (Figs 6 and 9).
Description.
Male (focal specimen: holotype). Carapace length 1.9; abdomen length 1.8. Palp (Figs 1–2) with bulb little rotated, embolus arising at about 140°; TmA thin and with only a hint of an elbow. RTA triangular. First leg with tarsus and metatarsus thicker than usual. Colour: Chelicerae pale, covered with erect white setae. Palp femur, patella, tibia with partially erect white setae, especially long prolaterally. Distal 3/4 of cymbium with fine dark hairs. Legs light to medium brown in alcohol, though darker in life. First leg metatarsus and tarsus dark, with extended dark scopula. Clypeus covered with cream coloured scales, with prominent row of long cream-coloured setae extending forward, forming a shelf (Fig. 7). Carapace dark brown with bronze scales. Abdomen similar to that of the juvenile in Fig. 12, reddish brown with broad paler basal band, two distinctive cream triangles centrally, and lateral cream bands made of paired crescents.Female (focal specimen: paratype, specimen from Estero Cerro Prieto, Figs 3–4). Carapace length 2.3; abdomen length 3.0. Epigynum with semicircular atria; central pocket broad posteriorly (Fig. 3). Colour: Chelicerae medium brown. Legs medium to pale brown, the first pair darkest. Clypeus covered with white scales, with (as in male) shelf of long white setae projecting forward (Fig. 10). Carapace and abdomen covered with cream-coloured scales except for orange-tan patches on abdomen. Central pale triangles (chevrons) on dorsum connected to lateral bands, as in Figs 8 and 10.
Additional material examined.
Two juveniles and 3 females from the type locality.
Natural history.
Found only in the negative tidal estuaries of Sonora, México. These unusual habitats have salt-tolerant plants (such as ) on soil that is constantly wet with salt water, as the tides enter then drain to cut stream-like channels (Figs 95–96). Fresh water is rarely available in this harsh desert. Although was found at Estero Morúa, in 2013 it was considerably more common at Estero Cerro Prieta. There, it was found either in retreats in the larger salt-tolerant plants along the edges of the channels, about 20-40 cm above the substrate, or on the mud/sand of the slopes of these channels after shaking the overhanging salt-tolerant plants. The courtship involves behaviours similar to those seen in other -group members, though of weak amplitude. A video of the courtship of male AZS13-7854 is available at https://youtu.be/JUkULLdOZ0w.
Figures 95–101.
Habitats of new and species. 95–97 Negative estuaries near Puerto Peñasco, Sonora 95, 96 Habitat of , type locality, Estero Cerro Prieto 97 Habitat of , type locality, Estero Morúa 98 Habitat of , type locality, Mt. Baldy, British Columbia 99–101 Tropical deciduous forest of Chamela, Jalisco at type locality for both and
99 Habitat of , type locality 100 Habitat of , type locality.
Male specimen JAL14-9837 in CNAN-IBUNAM, with data: México: Jalisco: Estación de Biología Chamela, 400-650 m on Calandria Trail, 19.5038 - 19.5045°N 105.0334 - 105.0344°W, 19 Feb. 2014, W. Maddison & H. Proctor WPM#14-034.
Males from the area of El Tuito, north of the type locality, have a continuous red patch in the centre of the clypeus (Fig. 21), instead of two vertical lines.12♂♂ 1♀ in UBC-SEM: México: Jalisco: El Tuito, Rancho Primavera, , 700 m elev., 3 March 2014, W. Maddison, WPM#14-047 (4♂♂); México: Jalisco: Sierra Manantlan, , 1550 m elev., 1 June 1998, W. Maddison et al., WPM#98-067 (7♂♂ 1♀). México: Jalisco: Estación de Biología Chamela, , 1-2 June 1998, W. Maddison et al., WPM#98-071 (1♂).Known from the tropical deciduous forests along the southern coast of Jalisco, México (Fig. 101), typically found on leaf litter or sticks on the ground that receives sun but is somewhat shaded (Fig. 99) — in contrast to the more open sunny ground on which lives nearby. The courtship involves the male standing at a distance from the female with first legs spread; he walks in bursts toward the female, sidling somewhat. On each burst forward, the front legs are flicked upward and the palps lowered to expose the face. A video of male courtship is available at https://youtu.be/mgXhB61u0mA.Maddison
sp. n.http://zoobank.org/1FCF2BA2-B4E8-4C5C-B384-7198F62E5D17Figs 26–37
cf. , from Lake Temagami, Ontario, male left palp, ventral view.
Note.
At the Royal Ontario Museum in 1978 I saw a male specimen of from Lake Temagami, Ontario, from whose label I recorded the collecting data “Ontario: Temagami. Island 1027. 24 June 1939. #5669”, although the museum reference notes indicated the date as 27 June 1937. It was notable for the brush of longer setae on the dorsal distal surface of the cymbium, and the twisted and tufted tarsus of the first leg. In both of these features it resembled the two described species (Peckham & Peckham, 1901) (from the southeastern U.S.) and Griswold, 1987 (from the southern Rocky Mountains of Wyoming, Colorado, and New Mexico), both of which are notable for the twisted and tufted tarsus and metatarsus of the first leg (Chamberlin and Ivie 1944, figure 210). I drew the palp (Fig. 71), which differs distinctly in rotation of the bulb from those two species (embolus arising at 225°, compared to 270° for and 290° for ), and thus represents a new species. I did not draw the cymbial brush or the first leg, and my memory does not retain their details except that there was a clear resemblance to in these ornaments. Recent attempts to locate the specimen at the museum have failed, and it may have been loaned for a project on (which was never completed) and not returned. In 1995 I travelled to the exact island in Lake Temagami on the label, but no specimens were found. However, the island was rock of perhaps 5 meters by 2 meters, unlikely to sustain any permanent population, and so either the specimen balPageBreaklooned in, or the label was incorrect. We are thus left with a biogeographically puzzling new species with no specimen on which to describe it. Two possible habitats might be productively searched: the rock outcrops of the Canadian Shield, or exposed sand of glacial deposits in Northern Ontario and Québec.cf. , from Lake Temagami, Ontario, male left palp, ventral view.Simon, 1876Walckenaer, 1802Currently 84 species are assigned to (World Spider Catalog, 2016). There are five subgenera: the nominate, three described by described by Logunov et al. (1999) and one by Prószyński (2016). These are:Simon 1876. Logunov et al. (1999) included three species in the nominate subgenus. The TmA is massive, much larger than the embolus. Logunov et al (1999) diagnosed
s. str. by “a raised epigynal median septum in females ... and the tibial apophysis appressed in the cymbial groove in males”.Logunov, Marusik & Rakov, 1999 (type species Kulczyński, 1895). Six species are assigned to . Their TmA is a more or less straight flat blade, wide especially at the tip, parallel to the embolus and as long as it. Logunov et al (1999) diagnosed by the RTA very short or absent, an apical spine on the TmA, a recessed epigynal atrium, and a one-chambered spermatheca.Logunov, Marusik & Rakov, 1999 (type species Simon, 1868). Twenty-three species are assigned to . They are compact-bodied (not elongate), with contrasting markings and a somewhat-ornamented male first leg, resembling in habitus to some extent the species group. The TmA is as long as the embolus but wider, with a complex pointed tip. Logunov et al (1999) diagnosed by the heavily sclerotized epigynal flaps and the subparallel tips of the embolus and the TmA.Logunov, Marusik & Rakov, 1999 (type species Logunov, 1995). Four species are assigned to , having a large complex TmA that PageBreakcurls distally at the tip, like a tongue. Logunov et al. (1999) diagnosed by the embolus and TmA perpendicularly orientated to each other and the peculiar elevated central pocket of the epigynum.Prószyński, 2016 (type species implied to be Sundevall 1833). has a long embolus and TmA, the latter much wider. Prószyński (2016) characterizes by the long needle-like embolus sheathed in the thick TmA, trapezoidal RTA, medial groove behind the central pocket of the epigynum, and spiral spermathecae.In all of these subgenera, the TmA is distinctly larger than the embolus. Many other species of are not yet assigned to a subgenus, including many African species whose TmAs are small or (apparently) absent.Although two Holarctic species are known from the Americas, and (Grube, 1861), the remaining species of New World form a distinctive group not known to occur in the Old World. The subgenus is here described to contain them.
. 72 Bulb of left palp, holotype of Emerton, 1925 73–74 Left palp, holotype of Lowrie & Gertsch, 1955 75–76 Left palp, male from Nova Scotia (DRM02.103) 77–78 Left palp, male from east-central Ontario (WPM#76-133) 79 Male from east-central Ontario (WPM#76-133) 80 Epigynum, ventral view, female from Nova Scotia (DRM02.103) 81 Epigynum, cleared, dorsal view, same female. DRM and WPM collecting codes are those indicated in specimen records.
http://zoobank.org/1F25FF98-0859-41E1-A72F-F56771B28AD3Figs 86–94Male (Figs 86, 87) in UBC-SEM, with data: Canada: British Columbia: Mt. Baldy. , 1180 m elev. 17 May 2013 W.Maddison & H. Proctor WPM#13-014.(3♂♂ 2♀♀). Same data as holotype (1♂ specimen NA13-6071 and 1♀ specimen NA13-6083 in UBC-SEM). Canada: British Columbia: W of Midway, along HWY 3, 3.0 km E of crossing of Kettle River with HWY 3, ca. , 2 May 1982, W. & D. Maddison, WPM#82-019 (1♂1♀ in CNC, 1♂ in AMNH).Named for the country of the type locality, in honour of the 150th anniversary of Canada’s confederation.A typical member of with striped markings, more contrasting in males than females. can be distinguished by the TmA being only slightly smaller than the embolus, diverging from the embolus initially, then curving distally to touch the tip of the embolus. has a similar palp, but its TmA is shorter and considerably narrower, only 1/4 to 1/3 the width of the embolus, and also is pressed against the embolus its entire length (Lowrie and Gertsch 1955, figure 17; holotype in AMNH examined).Male (focal specimen: holotype). Carapace length 2.0; abdomen length 2.1. Structure of body typical for . Embolus a short pointed blade, accompanied by a TmA of almost the same size, which opposes the embolus like a thumb against a forefinger (Fig. 86). The RTA is broad but pointed (Fig. 87). The cymbial lobe is small but distinct, projecting toward the retrolateral (e.g., visible behind the RTA in Fig. 86). Colour: Black except for paler femur, patella and tibia of palp, and coxae and trochanters of legs. Body with longitudinal stripes of white scales (Figs 90–91).Female (focal specimen: paratype, specimen NA13-6083, Figs 88, 89, 93, 94). Carapace length 2.4; abdomen length 2.7. Structure of body typical for . Central pocket not on a raised sclerotized mound (it is in ). Openings posterior to central pocket (Fig. 86), as in (Lowrie and Gertsch 1955, figure 29). Colour paler than male, with more distinct chevrons on the abdomen. Clypeus white except black patches below AME (Fig. 93).U.S.A: Montana: Glacier Co., 1.3 mi SE of intersection of HWY U.S. 89, Cutbank River & HWY 445, 24 May 1977, D., W., L., & R. Maddison, WPM#77-099 (12 ♂♂ 7♀♀ 4 juveniles in UBC-SEM).Collected at fairly high elevation on open ground with scattered small rocks, sticks and sparse vegetation (Fig. 98).Habitats of new and species. 95–97 Negative estuaries near Puerto Peñasco, Sonora 95, 96 Habitat of , type locality, Estero Cerro Prieto 97 Habitat of , type locality, Estero Morúa 98 Habitat of , type locality, Mt. Baldy, British Columbia 99–101 Tropical deciduous forest of Chamela, Jalisco at type locality for both and
99 Habitat of , type locality 100 Habitat of , type locality.Habitat of , Arizona, Amateur Astronomy Vista, .
Authors: Daniel B Zurek; Thomas W Cronin; Lisa A Taylor; Kevin Byrne; Mara L G Sullivan; Nathan I Morehouse Journal: Curr Biol Date: 2015-05-18 Impact factor: 10.834