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Fig Pollinators At Palomar College 3
© W.P. Armstrong Updated 11 April 2025

Tentative Hypotheses For New Fig Wasp Discoveries On Two Ficus Species At Palomar College: Afghan Fig (Ficus johannis ssp. afghanistanica) and Ficus socotrana (F. vasta). These observations & wasp collections made by Tony Rangel & Robert Kurner, April 2025.

Afghan Fig (Ficus afghanistanica)

Afghan Fig (Ficus johannis ssp. afghanistanica). Native to mountainous regions of N. India, W. Iran, & Afghanistan. According to peer-reviewed fig references, this species is closely related to F. carica & is pollinated by the same symbiotic wasp species (Blastophaga psenes). Like F. carica, it is apparently gynodioecious with female trees bearing syconia with only long-style female flowers and "male" trees bearing syconia with short-style female flowers and pollen-bearing anthers (male flowers). It also produces male & female fig wasps. Because the mature syconia appear to be "male", our tree would technically be classified as "male." In my opinion the wasps resemble the pollinator of F. carica (Blastophaga psenes). We certainly have a producer of this wasp species on campus: My 'Vista Caprifig' (Ficus carica). In fact, it superficially resembles F. pseudocarica, another taxonomic controversy in fig biology. At my home in Twin Oaks Valley the 'Vista Caprifig' is a super pollinator resulting in numerous fig seedlings at my home & neighborhood. This is why a professor at Cal State Northridge & fig grower wanted cuttings from my 'Vista Caprifig' because pollinated fig cultivars have superior tasting figs! That is also why bags of wasp-bearing caprifig syconia are placed in Calimyrna fig orchards of the Central Valley. In fact, caprifigs are grown on these farms for production of wasp-bearing syconia. [I have visited these orchards.]

  Calimyrna Fig In Central Valley  

Afghan Fig (Ficus johannis ssp. afghanistanica) at Palomar College. Although the leaf shapes can be variable, they often have deep lobes reminiscent of the shape of snow flakes.

Afghan Fig (Ficus johannis ssp. afghanistanica) at Palomar College. The presence of fig wasps and male flowers indicate that this tree is a "male" of a gynodioecious species. In Ficus carica male trees are called caprifigs. The prefix "capri" refers to goat because the inedible fig syconia were fed to goats instead of people. (Unless, of course, you like a mouthful of pollen and fig wasps!)

Afghan Fig (Ficus johannis ssp. afghanistanica) at Palomar College. This interesting sample was stuck to the inside of collection vial.

These sure look like male & female Blastophaga psenes wasps to me! They were in vial with syconia from Ficus johannis ssp. afghanistanica.


Ficus carica-F. palmata Complex & Possible Origin Of Ficus carica

Of all the different hypotheses regarding the origin of the Ficus carica-palmata complex, including F. pseudocarica, F. johannis & F. afghanistanica, there is one proposed by Louis Trabut a century ago that seems very plausible today. Trabut's article "Sur Les Origines du Figuier" appeared in the French journal Revue de Botanique Appliquee, Volume 2: 393-396 (1922). It is summarized here by Ali Sarkhosh, et al. in The Fig: Botany, Production and Uses, CAB International Publishers, one hundred years later (2022): Ficus carica may be a synthetic species derived from several wild species through natural hybridization. It is found near other closely related species or subspecies, such as F. persica, F. virgata and F. johannis; grown in southwestern Persia, Mesopotamia and Arabia. Trabut (1922) also mentions F. forskalaei & F. morifolia from Arabia and Abyssinia. Natural hybridization occurs between all these species, the resulting seedlings being exceedingly variable. F. palmata, though generally regarded as the Indian form of F. carica, is widely distributed from northern India to Afghanistan, Arabia, Egypt and Abyssinia. All the figs in this group are dioecious (gynodioecious) with female trees & male caprifigs in the populations. Throughout this vast area of the Middle East, they all have the same symbiotic pollinator wasp Blastophaga psenes. As noted in above paragraphs, misidentification between Ficus carica and F. palmata is common in herbaria and botanical gardens, and there is continuous variation between species in this complex.

Leaf variation from a single branch on 'Vista Caprifig' in Twin Oaks Valley: Upper & lower leaf surfaces from 2 leaves are shown. Based on leaf shape, they could have come from practically any tree in the Ficus carica-palmata complex. Leaf characteristics in dichotomous keys to species in this complex are not very useful.
Most of the fig "species" in the Ficus carica-palmata complex are listed as synonyms or
subspecies of each other in Kew Plants Of The World Online and World Flora Online

Treatment Of 8 Species Of Ficus In Carica-Palmata Complex
Kew Plants Of The World Online & World Flora Online (WFO)

Note: Ficus carica L. is listed as an accepted species in both Kew & WFO.
  Did it evolve from following wild species complex through natural hybridization?
  

8 Species in Carica -
Palmata Complex
Kew Plants Of The
World Online
World Flora Online (WFO)
 F. afghanistanica Warb. 
syn. F. johannis ssp.
  afghanistanica (Warb.) Browicz
  
syn. F. johannis ssp.
  afghanistanica (Warb.) Browicz
  
 F. forskalaei Vahl. 
syn. F. palmata ssp. palmata
syn. F. palmata Forssk.
F. johannis Boiss.
= F. johannis Boiss.
= F. johannis Boiss.
F. morifolia Forssk.
syn. F. palmata ssp. palmata
syn. F. palmata Forssk.
F. palmata Forssk.
= F. palmata Forssk.
= F. palmata Forssk.
F. persica Boiss.
syn. F. johannis ssp. johannis
syn. F. johannis Boiss
F. pseudocarica Miq.
syn. F. palmata ssp.
virgata Browicz
syn. F. palmata ssp.
virgata (Roxb.) Browicz
F. virgata Roxb.
syn. F. palmata ssp.
virgata Browicz
syn. F. palmata ssp.
virgata (Roxb.) Browicz

Author names should be listed for accurate synonym comparisons. For example,
F. palmata Rosb. is a synonym of a different species F. grossularioides Burm.f.

Ira Condit was well aware of Trabut's hypothesis for the origin of Ficus carica. In chapter 3 of his book The Fig (1947), Condit discusses natural hybridization between all the species in the F. carica-F. palmata complex resulting in exceedingly variable seedlings: "Trabut hoped to be able to remake this synthetic species [Ficus carica] by bringing together all the related forms and by allowing them to interpollinate. He was able, however, to secure only F. palmata and F. pseudocarica. In California as in Algeria these two species have long been acclimatized and are being utilized in certain lines of plant breeding."

Regarding all these synonymous or very closely related species in the Ficus carica-palmata complex, there is one undeniable fact. The oldest officially named species is Ficus carica L., named by Linnaeus & published in Species Plantarum in 1753. According to the International Codes of Botanical Nomenclature, Rule (Law) of Priority, this is a correct name that should be used!

  Aradhya et al. Cladogram Comparing DNA Of Ficus carica Varieties (PDF)  


Socotran Fig (Ficus socotrana = F. vasta)

Subgenus Urostigma Section Conosycea

1.   Indian Laurel Fig (Ficus microcarpa)

Subgenus Urostigma Section Malvanthera

2.   Rustyleaf Fig (Ficus rubiginosa)
3.   Moreton Bay Fig (Ficus macrophylla)
4.   Banana Fig (Ficus pleurocarpa)

Subgenus Urostigma Section Galaglychia

5.   Socotran Fig (Ficus vasta)

Monoecious fig species at Palomar College with fig wasps in their syconia. Some of these are tentative examples of host sharing. This list may be increasing as syconia of additional fig species on campus are carefully examined. Note: the wasp-bearing gynodioecious speces on campus (F. carica) and F. johannis ssp. afghanistanica belong to the subgenus Ficus, section Ficus and are pollinated by Blastophaga psenes.

Socotran fig (Ficus socotrana = F. vasta). Native to NE Tropical Africa to N. Kenya, Arabian Peninsula, Ethiopia & Yemen. [Including Socotra Island off the coast of Yemen.] The native pollinators are Ceratosolen arabicus & Elizabethiella socotrensis. The wasps I observed are definitely not Ceratosolen. It is doubtful that Elizabethiella reached Palomar College. In addition, the original name of holotype was Blastophaga socotrensis. It is interesting to note that Blastophaga psenes has also been reported from Yemen.

One additional comment about host sharing. Fig biologists originally thought each species of fig had its own unique symbiotic pollinator wasp; however, peer-reviewed publications during the past few decades have clearly shown that host sharing and pollinator sharing are a lot more common than previously thought. I have included the following section from my Pollinators 1 about this subject.

Host Sharing & Pollinator Sharing

Host Sharing: 2 or more pollinator wasp species per host.
Pollinator Sharing: 2 or more fig host species per pollinator.

Fig & Fig Wasp Pollination May Be Even More Complicated

When I began studying figs (Ficus) with horticulturist Steven Disparti over 30 years ago, I thought each species of fig had it own unique, symbiotic pollinator. This may still be true for some of the 876 species of Ficus; however, multiple pollinators and evidence from host & pollinator sharing data have complicated the fig & fig wasp story. The relationships undoubtedly follow kinships within subgenera & sections depicted in enormous Ficus DNA cladograms (evolutionary trees). There are 4 pollinated Australian fig species on campus in subgenus Urostigma and at least 3 verified species of Australian fig wasps: (12 possible interactions). If you exclude Section Conosycea & wasp Eupristina, and only consider the more closely related Section Malvanthera & 2 Pleistodontes wasp species on campus, there would be 7 possible interactions. Another complicating factor introduced into seedling diversity is possible hybridization between fig species pollinated by multiple wasp species each coming from at least 2 different host fig species. In fact, some authorities have speculated that host-sharing and host-switching by fig wasps may have lead to the enormous number of fig species. In host-sharing multiple fig species share the same pollinator wasp, while in host-switching a pollinator wasp evolves to use a different fig species as its primary host.

    Carlos A Machado, Nancy Robbins, M Thomas P Gilbert, & Edward Allen Herre. 2005. "Critical review of host specificity and its coevolutionary implications in the fig/fig-wasp mutualism." Proc Natl Acad Sci U S A. 2005 Apr 25; 102 (Suppl 1):6558–6565. doi: 10.1073/pnas.0501840102

    Gang Wang, et al. 2023. “Genomic evidence of prevalent hybridization throughout the evolutionary history of the fig-wasp pollination mutualism.” https://doi.org/10.1038/s41467-021-20957-3

    Rasplus, Jean-Yves. 1996. The Biodiversity of African Plants. "The One-to-One Species Specificity of the Ficus-Agaoninae Mutualism: How Casual?" Frech National Institute for Agriculture, Food, and Environment (INRAE). doi: 10.1007/978-94-009-0285-5_78

Socotran Fig (Ficus socotrana = F. vasta) at Palomar College.

Socotran Fig (Ficus socotrana = F. vasta) at Palomar College. I originally thought these female wasps were Blastophaga psenes from nearby caprifig (F. carica), but the head shape and ventral side did not match. The perfect fit was Pleistodontes imperialis, native pollinator of the rustyleaf fig (Ficus rubiginosa). [See Next Image/]

Mystery wasps in syconia of Socotran Fig (Ficus socotrana = F. vasta) at Palomar College. Based on image comparisons of the shape and ventral side of head, Elisabethiella socotrensis and Blastophaga psenes can be ruled out. The perfect fit appears to be Pleistodontes imperialis, native pollinator of the rustyleaf fig (Ficus rubiginosa), a tree well represented at Palomar College.

The 2 divergent rows of ridges are apparently called ventral lamellae (I think). They are apparently useful in identification of fig wasp species. Several online references say they may help the female move or transport pollen within the syconium. This species definitely packs pollen into her pouches (corbiculae) on the underside of her thorax.

Another newly discovered fig on campus (Ficus obliqua) also has Pleistodontes imperialis in its syconia! The leaves are smaller than F. rubiginosa. The syconia are slightly smaller.


Wasps at top of image will easily slip through
the "eye" of an ordinary sewing needle.

Summary and Conclusions

The 'Vista Caprifig' (Ficus carica or possibly F. pseudocarica) produces 3 crops of syconia per year, each crop releasing gravid (pregnant) pollinator wasps that perpetuate wasps by laying eggs in receptive syconia of following crop. (1) Profichi crop in late spring, (2) Mammoni crop in summer, & overwintering Mamme crop in the fall. The "Mamme" crop releases wasps the following year into receptive "Profichi" crop that mature & release wasps in late spring.

In the case of Afghan fig Ficus johannis ssp.afghanistanica, the presence of male and female wasps plus pollen-bearing anthers (male flowers) indicate that they emerged from short-style female flowers within syconium. I.e. this syconium must have been previously pollinated and eggs were laid by pollinator, presumably Blastophaga psenes. The timing of late March-April is too early for release of pollen-laden female wasps from 'Vista Caprifig' "profichi crop" on campus this spring. Maybe wasps came from overwitering "mamme" crop that may not have carried pollen. If so, the Afghan fig syconia would produce wasps but no viable seeds. Whether Afghan fig produces viable seeds remains to be seen.

Identity of wasps in monoecious fig (Ficus vasta) has been solved. They are Pleistodontes imperialis from rustyleaf fig (F. rubiginosa), a species of monoecious fig. A nearby monoecious fig, tentatively determined as Ficus obliqua also has syconia with P. imperialis. These appear to be more cases of "pollinator sharing" (2 or more fig host species per pollinator) at Palomar College. "Host Sharing" is 2 or more pollinator wasp species per host.

Pleistodontes imperialis in Syconia of Multiple Fig Species
(Pollinator Sharing: 2 or more fig host species per pollinator)


Kauai:

Subgenus Urostigma:

Ficus rubiginosa--Section Malvanthera

Ficus watkinsiania--Section Malvanthera

Ficus macrocarpa--Section Malvanthera (Pleistodontes froggatii)

Ficus rubra--Section Galoglychia Subsection Platyphyllae

Jared Bernard, et al. 2020. "New Species Assemblages Disrupt Obligatory Mutualisms Between Figs and Their Pollinators." Front. Ecol. Evol., 18 November 2020 Sec. Coevolution. Volume 8 - 2020 | https://doi.org/10.3389/fevo.2020.564653


Palomar College:

Subgenus Urostigma:

Ficus rubiginosa--Section Malvanthera Subsection Platypodeae

Ficus pleurocarpa? Section Malvanthera (Presumably P. imperialis)

Ficus macrophylla? Section Malvanthera (P. froggatii or imperialis?)

Ficus cf. obliqua--Section Malvanthera Subsection Platypodeae

Ficus vasta--Section Galoglychia Subsection Platyphyllae


Europe:

Subgenus Urostigma:

Ficus macrophylla columnaris--Section Malvanthera

Pleistodontes cf. imperialis (Following reference used cf.)

Manlio Speciale, Giuliano Cerasa & Gabriella Lo Verde. 2015. "First Record in Europe of Seedlings of Ficus macrophylla F. columnaris (Moraceae) and of Its Pollinating Wasp Pleistodontes cf. imperialis (Chalcidoidea Agaonidae)." Naturalista Sicil., S. IV, XXXIX (2), pp. 399-406.

Note: Sydney S. Saunders, the original author of Pleistodontes imperialis cited the host tree as Ficus macrophylla in 1882 (Entomologist's Monthly Magazine, Volume 19).