Short-legged daddy longleg spiders: evolution of the enigmatic Ninetinae
2019-2025, funded by the German Research Foundation, DFG
 
The daddy longlegs spider subfamily Ninetinae was previously composed of no more than 31 described species. Most Ninetinae are small to tiny cryptic ground-dwellers and restricted to arid habitats in the New World, Africa, and the Arabian Peninsula. As a result, Ninetinae have been poorly represented in collections, fresh material for molecular work was barely available, and almost nothing had been known about their biology.


Ninetinae (above), and ‘typical’ Pholcidae of other subfamilies (below).

At the same time, preliminary data had shown that Ninetinae deserve special attention for several reasons: (1) they seemed to be ‘basal’ within Pholcidae and thus crucial for reconstructing the evolutionary history of the family; (2) they differ from other Pholcidae in many ways, including morphology (e.g. leg length, sperm morphology), biology (e.g., web building, female sperm storage), and ecology (e.g., arid habitats). However, their exact phylogenetic position could not be resolved in a Sanger-based six-gene analysis; only a single species had been studied with respect to sperm morphology; and all we knew about Ninetinae biology came from anecdotal field observations.


Typical habitats of Ninetinae (left: Brazil, right: Oman). In stark contrast to most other pholcids, Ninetinae tolerate extreme aridity.

This project aimed to unravel the evolution of these enigmatic spiders in a large collaborative effort. We combined the expertise of a team of ten people in five countries and apply state of the art methodology in three interconnected fields: systematics and evolution, biogeography, and evolutionary morphology and biology.


  • We collected Ninetinae at nine expeditions to nine countries, resulting in ~3320 adult specimens, more than all previous collections worldwide taken together.
  • We generated a UCE dataset with a strong representation of Ninetinae, and combined these data with previously published and new Sanger sequences into the most comprehensive phylogeny in arachnology (>1000 species; 96% of all known pholcid genera).
  • We documented strong sampling biases, a phylogenetically conserved environmental niche for the subfamily, and used distribution modelling to predict occurrences of Ninetinae in several poorly sampled regions.
  • We analyzed in detail the ultrastructure of numerous species, added to two of the largest arachnological datasets on sperm ultrastructure and karyology, generated first data about internal anatomy, and studied for the first time the sexual biology of a Ninetinae spider.
As of early 2025, revisions of most genera have been completed. The last revision (Kambiwa) and the molecular phylogeny are still in work.

Huber BA, Meng G. 2025. Like grains of sand: Ninetis spiders on the Arabian Peninsula (Araneae: Pholcidae). Zootaxa 5563: 290-335. https://doi.org/10.11646/zootaxa.5563.1.19

Huber BA, Meng G, Král J, Ávila Herrera IM, Carvalho LS. 2024. Diamonds in the rough: Ibotyporanga (Araneae, Pholcidae) spiders in semi-arid Neotropical environments. European Journal of Taxonomy 963: 1-169. https://doi.org/10.5852/ejt.2024.963.2687

Huber BA, Meng G, Dederichs TM, Michalik P, Forman M, Král J. 2024. Castaways: the Leeward Antilles endemic spider genus Papiamenta (Araneae: Pholcidae). Invertebrate Systematics 38: IS23052. https://doi.org/10.1071/IS23052

Huber BA, Meng G, Cabra García J, Carvalho LS. 2024. Thriving in dry conditions: on the Neotropical spider genus Galapa (Araneae: Pholcidae). Zootaxa 5419: 301-347. https://doi.org/10.11646/zootaxa.5419.3.1

Huber BA, Meng G, Král J, Ávila Herrera IM, Izquierdo MA. 2023. Revision of the South American Ninetinae genus Guaranita (Araneae, Pholcidae). European Journal of Taxonomy 900: 32-80. https://doi.org/10.5852/ejt.2023.900.2301

Huber BA, Meng G, A Váldez-Mondragón, Král J, Ávila Herrera IM, Carvalho LS. 2023. Short-legged daddy-long-leg spiders in North America: the genera Pholcophora and Tolteca (Araneae, Pholcidae). European Journal of Taxonomy 880: 1-89. https://doi.org/10.5852/ejt.2023.880.2173

Huber BA, Meng G, Král J, Ávila Herrera IM, Izquierdo MA, Carvalho LS. 2023. High and dry: integrative taxonomy of the Andean spider genus Nerudia (Araneae: Pholcidae). Zoological Journal of the Linnean Society 198: 534-591. https://doi.org/10.1093/zoolinnean/zlac100