Attulus (Attulus) floricola (C.L. Koch, 1837)

Maddison, Wayne P., Maddison, David R., Derkarabetian, Shahan & Hedin, Marshal, 2020, Sitticine jumping spiders: phylogeny, classification, and chromosomes (Araneae, Salticidae, Sitticini), ZooKeys 925, pp. 1-54 : 1

Attulus (Attulus) floricola (C.L. Koch, 1837) Figures 33-35 , 49-53

Euophrys floricola C. L. Koch, 1837.

Attus palustris Peckham & Peckham, 1883 (specimens in MCZ labelled as types, examined, but see below).

Attus morosus Banks, 1895 (synonymized by Prószyński 1980; confirmed here by examination of holotype female in MCZ from Olympia, Washington).

Remarks.

A widespread Holarctic species often found in retreats in dry flower heads in wetter areas such as marshes, A. floricola is distinctive for the sharp white lines around the eyes of males, forming an apparent mask (Fig. 34 ). Attulus floricola has often been confused in the past with its close relatives, but the distinctions have been clarified considerably by Prószyński (1980) and Logunov and Kronestedt (1997).

We treat the North American populations as full floricola , not a distinct subspecies. While Nearctic populations were long recognized as a separate species palustris , Prószyński (1980) suggested they are conspecific with the Eurasian populations. He maintained them as a distinct subspecies, but he expressed doubt as to whether even that distinction was warranted. We concur with his skepticism. If any consistent differences exist between the continents, they are no more visible than any differences that might exist between the Eurasian and North American populations of other species for which we don’t recognize subspecies such as Sittisax ranieri , Attulus cutleri , Dendryphantes nigromaculatus (Keyserling, 1885), Pellenes ignifrons (Grube, 1861), and Pellenes lapponicus (Sundevall, 1833).

The results of our COI analysis of Palearctic and Nearctic floricola group (Fig. 104 ) show all floricola to be close on the gene tree, with the New World specimens in two clades (not clearly related to one another) and the German specimens in a third clade. This suggests that A. floricola is not cleanly or deeply divided between the Nearctic and Palearctic. The molecular and morphological evidence leads us to fully synonymize palustris into floricola .

Within North America, the characterization of A. floricola has been muddied by confusion with a second species, A. sylvestris . Attulus sylvestris , long synonymized with palustris , is a distinctively different species. Attulus floricola is larger-bodied, has a much more contrasting colour pattern, and longer legs. Attulus floricola has a different angle of the spermaphore loop (subtle but consistent; Fig. 49 vs. Fig. 54 ), and in females the darkness of the spermathecal lobe is visible through the anteriormost portion of the epigynal atrium (Fig. 50 vs. Fig. 55 ). Attulus sylvestris has genitalia more similar to those of the Eurasian A. caricis , A. rupicola , and A. inexpectus , as noted below. The synonymy of sylvestris with palustris was originally proposed by Peckham and Peckham (1909), after which Kaston (1948) may have stirred confusion by choosing to illustrate palustris using Emerton’s (1891) figure of sylvestris .

A more serious confusion apparently occurred with the labelling of type specimens of Attus palustris . The description by Peckham and Peckham (1883) refers without doubt to the common white-striped species long known as Sitticus palustris (Fig. 34 ): males dark brown, reddish toward eyes, marked with white lines, including those around the eyes, and palp with some white hairs on several segments of the palp. As well, the habitat suggested by the name " palustris " is marsh or swamp, more typical for A. floricola than A. sylvestris . However, the specimens labelled as the types of Attus palustris in the MCZ are clearly specimens of the less common dusty brown species (i.e., Emerton’s sylvestris , Fig. 32 ). These specimens, we argue, are mislabelled: they do not match the Peckhams’ description, and thus are not the type specimens of A. palustris . That the Peckhams viewed the white-striped form as typical palustris can be judged not only from their 1883 description, but also from their implicitly distinguishing two forms in their 1909 statement "Mr. Emerton agrees with us that the form which he described as sylvestris is a variety of palustris , with the leg a little shorter and stouter." The label of the holotype does not appear to be in the handwriting of either George or Elizabeth Peckham, and it is possible that these “types” were so labelled after 1883.

At stake is not the name used for the common white-striped species (which would be floricola regardless), but the name for the uncommon dusty brown species, which would be palustris were we to accept these specimens as its types. However, as argued above, they are not the types. We therefore treat palustris as a synonym of floricola , and sylvestris as the name for the dusty brown species. To settle the mislabelling properly, a male specimen of the white-striped species from Wisconsin (the type locality) should be designated as the neotype or lectotype of palustris . We have not yet done so as we await reexamination of the full Peckham collection in case specimens can be located that might be identifiable as from the true type series.

Material examined.

Canada: British Columbia: Richmond (2 females), 49.66, -114.73 (1 female), 49.45, -115.08 (3 males, 6 females); Alberta: 52.46, -113.94 (1 male); Ontario: Richmond (2 males, 1 female), Gravenhurst (3 males), Port Cunnington (1 female); Dwight (2 males, 5 females), Batchawana Bay (1 female), Woodstock (3 females), 46.9300, -79.7268 (2 males, 1 female), 42.53, -80.12 (1 female), 43.2626, -80.5636 (1 male), 49.0852, -81.3237 (1 female); Quebec: Touraine (1 male); Nova ScotiA: 44.4318, -64.6075 (1 male); U.S.A.: Washington: 46.43, -123.86 (2 males); Colorado: Jackson Lake State Rec. Area (1 male); Nebraska: 41.88, -103.09 (1 female).

Figures

Figure 104. Relationships among Attulus floricola mitochondrial COI sequences in the context of the floricola group. Specimens in bold had their relationships constrained by the UCE phylogeny of Fig. 48; not shown are the relationships outside the floricola group, which are fixed to match the UCE phylogeny. The placement of non-bold specimens on this constrained skeletal tree was inferred by maximum likelihood (RAxML, codon positions as separate partitions).

Figures 31 - 38. Attulus subgenus Attulus, continued (floricola group) 31, 32 Attulus sylvestris: 31 male, Ontario, Ottawa 32 male, Maryland, Dorchester Co 33 - 35 A. floricola: 33 male, Ontario, Port Cunnington 34 male, Ontario (46.9300, - 79.7268) 35 male, Ontario, Gravenhurst 36 - 38 A. inexpectus: 36, 37 male, Tuva (50.6690, 92.9844) 38 female, Tuva (51.316, 94.495). For additional images of the floricola group, see Figs 49 - 58.

Figures 49 - 68. Sitticines of Canada: Attulus subgenus Attulus (for A. ammophilus, see Figs 69 - 73) 49 - 53 Attulus floricola: 49 palp (Ontario, Gravenhurst) 50, 51 ventral view of epigyne, dorsal view of cleared vulva (Ontario, Gravenhurst) 52 male (Ontario, 46.9300, - 79.7268) 53 female (Ontario, 46.9300, - 79.7268) 54 - 58 Attulus sylvestris: 54 palp (Ontario, Ottawa) 55, 56 ventral view of epigyne, dorsal view of cleared vulva (Ontario, Ottawa) 57 male (California, 36.3646, - 121.5544) 58 female (Ontario, 42.55, - 80.13) 59 - 63 Attulus striatus: 59 palp (Ontario, 45.1453, - 75.8467) 60, 61 ventral view of epigyne, dorsal view of cleared vulva (Ontario, 45.1453, - 75.8467) 62 male (Ontario, 45.1453, - 75.8467) 63 female (New Hampshire, Ponemah Bog) 64 - 68 Attulus cutleri: 64 palp (Northwest Territories, Wrigley) 65, 66 ventral view of epigyne, dorsal view of cleared vulva (Northwest Territories, Wrigley) 67 male (Northwest Territories, Inuvik) 68 female (Yukon, 67.57, - 139.67). For habitus of other Attulus species, see Figs 15 - 38.