Diphya, NICOLET, 1849

Álvarez-Padilla, Fernando & Hormiga, Gustavo, 2011, Morphological and phylogenetic atlas of the orb-weaving spider family Tetragnathidae (Araneae: Araneoidea), Zoological Journal of the Linnean Society 162 (4), pp. 713-879 : 756-761

publication ID	https://doi.org/ 10.1111/j.1096-3642.2011.00692.x
persistent identifier	https://treatment.plazi.org/id/7D5E87AD-C042-5536-FC34-4C97D2C4FA0D
treatment provided by	Valdenar
scientific name	Diphya
status

Treatment

DIPHYA NICOLET, 1849 View in CoL View at ENA ( FIGS 26–30 View Figure 26 View Figure 27 View Figure 28 View Figure 29 View Figure 30 )

Type species: Diphya macrophthalma Nicolet, 1849 . The holotype of D. macrophthalma is a female specimen from Valdivia ( Chile), deposited at the Museum National d’Histoire Naturelle, Paris (examined).

Diagnosis: Diphya species can be distinguished from all other tetragnathid genera by the following combination of characters: AME considerably smaller and PLE on separate tubercles ( Fig. 27E View Figure 27 ); secondary eyes without reflective tapetum; legs I and II armed with a mesal row of macrosetae; epigynum a flat sclerotized plate with the copulatory openings ventrally orientated ( Fig. 28A, B View Figure 28 ); spermathecae well sclerotized ( Figs 28C View Figure 28 , 30C View Figure 30 ); palpal tibia with a distal apophysis ( Fig. 29F View Figure 29 ); cymbium only with the cymbial ectomedian process ( Fig. 29D View Figure 29 ); paracymbium with an anterior apophysis, distally swollen ( Fig. 29C View Figure 29 ); spherical tegulum ( Fig. 29A View Figure 29 ). Conductor of membranous appearance but with the margins well sclerotized, coiled and firmly attached to the tegulum centre ( Fig. 29G View Figure 29 ). Embolus lamelliform and coiled, opposite to conductor ( Fig. 29E View Figure 29 ).

Description: Female: body length c. 0.5 mm. Transverse and shallow cephalothorax thoracic fovea ( Fig. 27A View Figure 27 ). Ocular area higher than carapace lateral margins ( Fig. 27E View Figure 27 ). Sternum as wide as long, with sculpted cuticle ( Fig. 27F View Figure 27 ). Cheliceral boss present, anterior cheliceral cuticle smooth ( Fig. 27D View Figure 27 ). Labium trapezoidal, wider than long and rebordered ( Fig. 27F View Figure 27 ). Anterior cheliceral cuticle as in clypeal area. Clypeus height less than one PLE diameter, AME considerably reduced. Abdomen oval lacking tubercles and silver guanine patches. Median tracheae not ramified, leaf-shaped apically ( Fig. 26A, E View Figure 26 ). ALS with c. 40 piriform spigots ( Fig. 26F View Figure 26 ). PMS anterior surface without aciniform spigots. PLS with fewer than 20 aciniform spigots arranged roughly in one line, aggregate spigots tips not embracing the flagelliform spigot apex ( Fig. 26D View Figure 26 ). Copulatory and fertilization ducts also well sclerotized and considerably shorter than half the spermatheca length ( Figs 28C View Figure 28 , 30C View Figure 30 ). Accessory gland openings immersed in individual pits, located away from the copulatory and fertilization ducts ( Fig. 28C, E View Figure 28 ).

Male: body length c. 2.8 mm. Somatic morphology as in female, except that the chelicerae are longer, with rugose cuticle on the anterior surface ( Fig. 27C View Figure 27 ). Epiandrous fusules concentrated in clusters, arranged in one transverse line ( Fig. 26G View Figure 26 ), with bases not immersed in pits. PLS triplet reduced to nubbins. Male palpal tibia as wide as long, with a hook-shaped apophysis on the ectodistal margin ( Fig. 29F View Figure 29 ). Palpal patella with one macroseta. Conductor membranous, with sclerotized edges at the base and apex, originating at the centre of the tegulum ( Fig. 30A View Figure 30 ). Sperm duct spiralled ( Fig. 30B View Figure 30 ).

Natural history: Diphya includes 11 species with a widespread geographical distribution that includes Argentina, Brazil, Chile, China, Japan, Korea, Madagascar, South Africa, and Taiwan. The type species is from Chile ( Nicolet, 1849; Simon, 1894; Tanikawa, 1995; Vellard, 1926; Zhu, Song & Zhang, 2003). Nothing is known about their biology and whether they build orb webs or not remains undocumented ( Namkung, 2003).

Taxonomy: Diphya has been revised by Tanikawa (1995), who described the first species from China, Japan, Korea, and Taiwan. Tanikawa (1995) revised and illustrated all known species except Diphya bicolor Vellard, 1926 (from Brazil) and Diphya simoni Kauri, 1950 (from South Africa). Our diagnosis and genus description is based on these studies plus additional specimens of an unnamed Diphya from Tanzania. The anatomy of D. spinifera has been studied in more detail than that of the other species; therefore, some characters in the description refer only to this species. Although the monophyly of Diphya remains untested by quantitative cladistic methods, similarity in the anatomy of all the species described and illustrated by Tanikawa (1995) suggests the monophyly of these taxa, except for the Malagasy species (unpubl. data). The present morphological and behavioural analysis proposed Diphya as sister to Tetragnathinae ( Fig. 143A View Figure 143 ); however, when molecular data are combined, Diphya and Chrysometa are recovered as sister taxa. In this last analysis there were no sequences for Diphya ; therefore its position as sister to Chrysometa is a result of the indirect optimization of morphological characters imposed by the congruence with the total evidence topology. We coded specimens of D. spinifera Tullgren, 1902 in the phylogenetic analysis.

DOLICHOGNATHA O. P.- CAMBRIDGE 1869

( FIGS 3E View Figure 3 , 31–35 View Figure 31 View Figure 32 View Figure 33 View Figure 34 View Figure 35 )

Type species: Dolichognatha nietneri O. P.- Cambridge 1869. The holotype of D. nietneri is a male specimen from Sri Lanka deposited in The Natural History Museum, London (O. P.- Cambridge, 1869; Levi, 1981).

Diagnosis: Dolichognatha species can be distinguished from all other tetragnathids by the following combination of characters: PME close together, when present, and smaller than the PLE ( Fig. 32A View Figure 32 ); clypeus height less than one AME diameter; chelicerae longer than the cymbial width, male chelicera longer, in some species longer than the carapace ( Fig. 32C, D View Figure 32 ); abdomen wider than long, with tubercles ( Fig. 35B View Figure 35 ); male palpal patella macroseta absent ( Fig. 32E View Figure 32 ); cymbial ectobasal process present ( Fig. 34A, B View Figure 34 ); metaine embolic apophysis present ( Figs 34E View Figure 34 , 35A View Figure 35 ).

Description: Female: body length from 2.6 to 4.0 mm. Transverse and shallow thoracic fovea, cephalic region considerably longer than the thoracic region ( Figs 32A View Figure 32 , 35B View Figure 35 ). Female sternum sculpted and longer than wide ( Fig. 32F View Figure 32 ). Labium trapezoidal, wider than long and rebordered. Ocular area higher than the carapace lateral margins ( Fig. 32E, G View Figure 32 ); lateral eyes on separated tubercles ( Fig. 32E, G View Figure 32 ). Cheliceral boss present, cuticle of anterior cheliceral surface rugose ( Fig. 32B–D View Figure 32 ). Abdomen with four pairs of caudal tubercles ( Fig. 35B View Figure 35 ), without silver guanine patches. Median tracheae considerably shorter than the lateral tracheal trunks, not ramified and with rounded tips ( Fig. 31A–D View Figure 31 ). Tracheal spiracle near the spinnerets, with fewer than four accessory glands on each side ( Fig. 31F View Figure 31 ). ALS with many piriform spigots. PMS anterior surface without aciniform spigots. PLS aggregate distal end apart from the flagelliform spigot ( Fig. 31E View Figure 31 ; Hormiga et al., 1995: fig. 24A–D). Epigynum well sclerotized, membranous in the middle, and slightly swollen. Copulatory openings as grooves orientated posteriorly ( Figs 33A, B View Figure 33 , 35D View Figure 35 ). Spermathecae spherical, well sclerotized, with one to two accessory glands ( Fig. 33C–E View Figure 33 ). Copulatory and fertilization ducts well sclerotized, running in parallel before entering the spermathecae ( Fig. 33D View Figure 33 ).

Male: body length from 2.6 to 3.2 mm. Somatic morphology as in female, except chelicerae, which are considerably longer ( Fig. 32E View Figure 32 ; Lise, 1993: figs 1–8). PLS triplet reduced to nubbins. Epiandrous fusules concentrated in two clusters, immersed in individual pits ( Fig. 31G View Figure 31 ). Male palpal tibia length approximately twice as long as wide. Conductor well sclerotized, fused to the tegulum, with a membranous apex. Tegulum larger than the subtegulum, cup-shaped ( Fig. 35A View Figure 35 ). Embolic metaine apophysis present, fused to embolus base, and longer than wide. Embolus short and tubular, similar in shape to the basal apophyses ( Figs 34E View Figure 34 , 35A, C View Figure 35 ). Sperm duct with two switchbacks ( Fig. 35C View Figure 35 ).

Natural history: Dolichognatha includes 27 species with a geographical distribution that includes the Old and New World tropics (O. P. Cambridge, 1869; Keyserling, 1883; Simon, 1895; Lessert, 1938; Marples, 1955; Tanikawa, 1991; Lise, 1993; Brescovit & de Cunha, 2001; Smith, 2008). The best documented species by far is Dolichognatha pentagona ( Hentz, 1850) , with a geographical distribution that extends from Venezuela to the southern USA ( Levi, 1981). Dolichognatha pentagona builds horizontal webs with more than 30 spirals and radii, sometimes with secondary radii and a closed hub. These webs are usually found between tree buttresses and roots or near the ground ( Fig. 3E View Figure 3 ). The web building behaviour of D. pentagona has been described by Eberhard (1982).

Taxonomy: Dolichognatha has never been revised and its monophyly has never been tested with a global taxon sample. A recent study included four Dolichognatha species from South-East Asia and Australia, and recovered this genus as monophyletic, supported by four synapomorphies: male versus female cheliceral size larger, palpal patellar macroseta absent, paracymbium secondary process procurved and metine embolic apophysis present ( Smith, 2008). We coded specimens of D. pentagona in the character matrix. Information available in the descriptions also includes the following species: Dolichognatha maturaca Lise, 1993 and the species included in Brescovit & de Cunha (2001) and Smith (2008). Two hypotheses have been proposed regarding the sister taxon of Dolichognatha : as either sister to Azilia ( Hormiga et al., 1995) or Metellina ( Álvarez-Padilla, 2007) . This latter sister group relationship is recovered in this study with the morphology and behavioural data set under all weighting criteria ( Fig. 143B View Figure 143 ). With the addition of DNA sequence data Dolichognatha was proposed as sister to a clade formed by Chrysometa and Diphya ( Fig. 144 View Figure 144 ).