Pholcid spiders: preliminary phylogeny and supporting characters
 
For illustrations of the characters supporting each node, klick on the node number or the taxon  name

Node
Taxon
Putative synapomorphies
1
Pholcidae
- Entire male palp modified (including proximal segments) (Huber 2000)
- Prominent paracymbium ('procursus') (Huber 2000)
- Male chelicerae sexually modified (Huber 2000)
- Clypeus about as high as chelicerae long (Huber 2000)
- Epigynum (not homologous to epigynum of Entelegynae) (Huber 2000)
- Only three trichobothria on walking leg tibiae (Huber 2000)
- Tarsal pseudosegments (Huber 2000)
- Arrangement of eyes (two triads separate from the anterior median eyes) (Huber 2000)
- ?Tarsus 4 comb hairs (Huber & Fleckenstein 2008)
- Egg sacs carried by females with their chelicerae
- ?Molecular data (Bruvu-Madaric et al. 2005; Dimitrov et al. 2013)
1a NN (all pholcids except Ninetinae) - ?Wide sternum
- ?Long legs (tibia 1 >2.5 x carapace width) and possibly correlated characters (retrolateral trichobothrium very proximal, many pseudosegments) (Huber 2000)
-
?Molecular data (12S, total evidence: Bruvo-Madaric et al. 2005)
1b NN (Arteminae + Modisiminae) - Molecular data (12S+CO1+16S+28S: Bruvo-Madaric et al. 2005; 28S: Astrin et al. 2007; Dimitrov et al. 2013)
1c NN (Smeringopinae + Pholcinae) - Tarsus 4 comb hairs over entire length of tarsus (Huber & Fleckenstein 2008)
-
Molecular data (12S+CO1+16S+28S: Bruvo-Madaric et al. 2005; CO1, 16S, 28S: Astrin et al. 2007; Dimitrov et al. 2013)
2
Ninetinae
- 'Ninetine tarsal organ' (small size, narrow orifice <35% of diameter) (Huber 2000)
- ?Short walking legs (tibia 1 <2.5x carapace width) and possibly correlated characters (retrolateral trichobothrium very distal, few pseudosegments) (Huber 2000)
- Compact ocular area with the anterior median eyes in a very high position (Huber 2011)
- ?Molecular data (Astrin et al. 2007; Dimitrov et al. 2013)
2a NN (Nerudia + Ninetis + Kambiwa) - Bulb with long spine-like projection (Huber 2000)
2b NN (Galapa + Guaranita) - Large dorsal flap on the procursus (Huber 2000)
2c
NN (Gertschiola + Ibotyporanga)
- Long procursus with characteristic proximal course (Huber 2000)
2d
NN (Tolteca +  Pholcophora + Papiamenta) - Simplified procursus (Huber 2000)
3 Arteminae - Procursus with dorsal apophysis and corresponding ventral pocket (Huber 2001)
-
Molecular data (CO1, 16S, 28S: Bruvo-Madarić et al. 2005; Astrin et al. 2007; Dimitrov et al. 2013)
3a
NN (all Arteminae except Artema) - Reduced epiandrous spigots (Huber 2000)
3b
NN (Aucana, Nita, Chisosa, Wugigarra, Tibetia, Holocneminus)
- Exposed tarsal organ (Huber 2000; Huber 2001; Huber & El Hennawy 2007; unpublished)
4 Modisiminae - Exposed tarsal organ (Huber 2000)
- Reduction of epiandrous spigots (Huber 2000)
-
Large distance between ALE and PME (Huber 2000)
- Molecular data (28S, total evidence:
Bruvo-Madaric et al. 2005; 16S+CO1, 28S: Astrin et al. 2007)
4a
NN (Modisiminae except Priscula) - Reduced ALS piriform spigots (Huber 2000, 2001)
- Male palpal coxa with retrolateral apophysis (Huber 2000)
- Molecular data (28S, total evidence: Bruvo-Madaric et al. 2005; 16S+CO1, 28S: Astrin et al. 2007)
4b
NN (Modisimus and potential close relatives) - Male palpal femur with pointed and upward projecting apophysis (Huber 1998, 2000)
4c
NN - Male femora and/or tibiae with vertical hairs in high density (Huber 2000)
4d
NN (Modisimus + Platnicknia)
- Molecular data (unpublished)
4e
NN (Aymaria + Ixchela) - ?Similar habitus (Huber 2000)
4f
NN (Coryssocnemis and potential close relatives)
- Male femur with bulge or apophysis directed towards proximal (or away from the femur at a right angle) (Huber 2000)
- ?Molecular data (28S, 12S: Bruvo-Madaric et al. 2005; CO1, 16S, 28S: Astrin et al. 2007)
4g
NN (Carapoia + Mesabolivar) - ?Molecular data (CO1, 16S, 28S: Astrin et al. 2007)
4h
NN (Canaima + Blancoa) - ?Similar habitus (Huber 2000)
5 Smeringopinae - ?Thoracic pit (Huber 2000; Huber 2012)
- Abdominal ventral pattern (three parallel dark lines (Huber 2012)
- Molecular data
(12S, CO1, 16S, 28S: Bruvo-Madaric et al. 2005; 28S: Astrin et al. 2007; Dimitrov et al. 2013)
-
?Silk puffs in webs (not obligatory) (Huber 2011; Huber 2012)
5a
NN (Smeringopinae except Smeringopus and Smeringopina) - Reduced ALS piriform spigots (Huber 2012)
- Reduced prolateral trichobothrium on tibia 1 (Huber 2012)
5b
NN (Stygopholcus + Hoplopholcus)
- Male cheliceral apophyses: two or more modified hairs each (Huber 2012)
5c
NN (Holocnemus + Crossopriza)
- Spotted legs (Huber 2012)
- ?Molecular data (28S, total evidence: Bruvo-Madaric et al. 2005)
5d
NN (Subsahara Smeringopinae: Smeringopus + Smeringopina)
- Two epiandrous spigots (Huber 2012, 2013)
- Reduced cheliceral stridulation (Huber 2012)
- Molecular data (Dimitrov et al. 2013)
6 Pholcinae - Male lateral (proximal) cheliceral apophyses (Huber 1995, 2000)
-
Tarsus 4 comb hairs in single row (Huber & Fleckenstein 2008)
- ?Molecular data (
12S, 28S, total evidence: Bruvo-Madaric et al. 2005; 16S+CO1, 28S: Astrin et al. 2007; Dimitrov et al. 2013)
6a
NN (Zatavua + Nyikoa) - Male palpal tarsus joints shifted to prolateral side (Huber 2007)
- ? Molecular data (Dimitrov et al. 2013)
6b
NN (Pholcinae except Zatavua and Nyikoa) - Male palpal trochanter with retrolateral apophysis
- ?Molecular data (12S, 28S, total evidence: Bruvo-Madaric et al. 2005; 16S+CO1, 28S: Astrin et al. 2007)
6c
NN (Spermophora and close relatives) - ?Genital bulb attached dorsally to cymbium
6d
NN (Buitinga and close relatives)
- Reduced ALS piriform spigots (Huber 2003a, b, 2005)
6e
NN (Buitinga + Spermophora Africa part)
- Molecular data (Dimitrov et al. 2013)
6f
NN (Quamtana and Pholcus plus close relatives) - Genital bulb with strong proximal apophysis
- ?Molecular data (28S, total evidence:
Bruvo-Madaric et al. 2005; Dimitrov et al. 2013)
6g
NN (Pholcus and close relatives) - Simplified tarsus 4 comb hairs (single row of tines) (Huber & Fleckenstein 2008)
-
Knob-like structure on epigynum (Huber 2011)
- ?Molecular data (28S, total evidence: Bruvo-Madaric et al. 2005; 16S+CO1, 28S: Astrin et al. 2007; Dimitrov et al. 2013)
6h
NN (Pholcus and close relatives except Sihala)
- Procursus with ventral projection ("knee") (Huber 2011)
- Genital bulb with additional apophysis (uncus) (Huber 2011)
6i
NN (Micropholcus + Pehrforsskalia + Leptopholcus + Micromerys) - Molecular data (Dimitrov et al. 2013; Huber et al. in press)
6j NN (Pehrforsskalia + Leptopholcus + Micromerys) - Male chelicerae with lateral apophyses very distally, no frontal apophyses (Huber 2011)
- Reduced epigynal knob (Huber 2011)
6k
NN (Leptopholcus + Micromerys) - Tip of male palpal trochanter apophysis with characteristic ridges (Huber 2011)
- Procursus ventral "knee" with transparent process (Huber 2011)
-
Reduced ALS piriform spigots (Huber 2001, 2011)