Arac 51 01 1

2023.
Journal of Arachnology 51:1–18
REVIEW
Fungus and fruit consumption by harvestmen and spiders (Opiliones, Araneae):

the vegetarian side of two predominantly predaceous arachnid groups
Martin Nyffeler1, Glauco Machado2, Arik Hartmann3 and Carl N. Keiser3: 1Department of Environmental Sciences,
University of Basel, CH–4056 Basel, Switzerland. E-mail: martin.nyffeler@unibas.ch 2LAGE do Departamento de
Ecologia, Instituto de Biociências, Universidade de São Paulo, 05508–090, São Paulo, Brazil; 3Department of Biology,
University of Florida, Gainesville, FL 32611, USA.
Abstract. In this review, we report on harvestmen and spiders feeding on fungi, fruits, and seeds. Fungivory in harvestmen
is widespread, with most reports referring to tropical species in the family Sclerosomatidae, which consume mainly small
forest mushrooms (families Marasmiaceae and Mycenaceae). In contrast, consumption of fungal material by spiders
apparently occurs only if airborne spores trapped in the viscid threads of orb-webs (e.g., Araneidae and Tetragnathidae) are
ingested along with old webs prior to the construction of new webs. Consumption of fruit pulp by harvestmen is also
widespread, with several records of Leiobunum spp. (Sclerosomatidae) feeding on Rubus spp. berries and other lipid-poor
fruits in the Holarctic region. In Neotropical forests, harvestmen in the families Cosmetidae and Gonyleptidae feed on lipid-
poor pulp of fallen fruits. Among spiders, we document several cases of synanthropic species opportunistically feeding on
fruit waste (e.g., pieces of banana, papaya, watermelon, or orange pulp) inside houses or disposed in yards. Only one case of a
spider feeding on a wild fruit in the field was found in our search. Finally, we report several cases of harvestmen and spiders
feeding on elaiosomes or arils (i.e., lipid-rich seed appendages). In conclusion, harvestmen consume mushrooms, fruit pulp,
seeds, and seed appendages more frequently than spiders probably because they are ‘‘solid food feeders’’, which means they
can ingest solid tissues by biting off small pieces. In turn, spiders are ‘‘fluid feeders’’ and feed on vegetable matter most
frequently in the form of fluids (e.g., nectar, stigmatic exudate, plant sap, and honey dew), rather than fungal or plant tissues.
Keywords: Aril, caloric value, diet, elaiosome, fruit waste, fungal spores, mushrooms, seeds.
https://doi.org/10.1636/JoA-S-22-015
TABLE OF CONTENTS
1. INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
2. METHODS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2.1 Data collection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2.2 Taxonomic comments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2.3 Chemical property information for fungi and fruits. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2.4 Assessment of the caloric value of fungi and fruits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3. RESULTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3.1 Fungivory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3.1.1 Feeding on mushroom pileus or sap . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3.1.2 Feeding on fungal spores and yeast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
3.2 Frugivory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
3.2.1 Feeding on fruit pulp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
3.2.2 Feeding on seeds and seed appendages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
4. DISCUSSION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
4.1 How common is fungivory and frugivory in harvestmen and spiders? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
4.2 Which taxa are engaged in fungivory and frugivory? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
4.3 What are the nutritional benefits of fungivory and frugivory?. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
4.4 Can harvestmen and spiders sustain on only fungal or fruit diets? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
5. CONCLUDING REMARKS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
1. INTRODUCTION Species in these orders exhibit an enormous diversity of lifestyles

and foraging strategies in terrestrial ecosystems (Machado et al.
Harvestmen (Opiliones) and spiders (Araneae) are two 2007; Foelix 2011; Nyffeler & Birkhofer 2017). Harvestmen have
speciose arachnid orders, which comprise together more than been conventionally described as predators or omnivores and
56,000 species (Kury et al. 2021; World Spider Catalog 2022). spiders as predators; both groups, however, depend largely on
1
2 JOURNAL OF ARACHNOLOGY
arthropods as food (Nentwig 1987; Wise 1993; Nyffeler et al. following the same search method as Nyffeler et al. (2017c).
1994; Acosta & Machado 2007). More recent studies expanded The combination of key words used in our search is presented
our understanding of the foraging behavior and diet of in Appendix 1 (see Supplemental Materials, online at https://
harvestmen and spiders, including observations of individuals doi.org/10.1636/JoA-S-22-015.s1). Additionally, we made a
feeding on small vertebrates (e.g., Castanho & Pinto-da-Rocha library search of books and scientific journals not included in
2005; Benson & Chartier 2010; Oliveira et al. 2010; Nyffeler et al. the electronic databases. Social media sites (e.g., BugGuide,
2017a; Nyffeler & Gibbons 2022a) and other unusual prey, such iNaturalist, Facebook, Flickr, Getty Images, Reddit, Yahoo,
as gastropods (Nyffeler & Symondson 2001), earthworms and YouTube) were also searched. Some of the bloggers who
(Nyffeler et al. 2017b), and eggs of invertebrates and vertebrates had posted photos and/or reports relevant to the review were
(Nyffeler & Gibbons 2022b). contacted by us to obtain additional information. For
In 2007, Acosta & Machado compiled a list of the various harvestmen, a total of 96 records have been found, 41 of
types of food items consumed by harvestmen, including reports which refer to consumption of fungal material, 39 to
of consumption of fruits, seeds, pollen, mushrooms, lichens, and consumption of fruit pulp, and 16 to consumption of seeds
algae. Almost ten years later, Nyffeler et al. (2016) published a or seed appendages (Tables 1–3). By contrast, a total of 28
review of plant-eating by spiders. These reviews bear witness that records relating to spiders have been found, 3 of which refer to
both harvestmen and spiders feed on a broad range of fungal consumption of fungal material, 12 to consumption of fruit
and/or plant materials in addition to their usual arthropod prey. pulp, and 13 to consumption of seeds or seed appendages
It must be said, however, that the number of reports on (Tables 1–3).
consumption of plant materials by harvestmen is very limited in 2.2 Taxonomic comments.—Nomenclature of harvestman
number (Acosta & Machado 2007). For spiders, the number of and spider taxa is based on the World Catalog of Opiliones
reports of consumption of plant materials is considerably higher, (http://wcolite.com/) and World Spider Catalog (https://wsc.
but most of them refer to cases in which nectar and pollen are nmbe.ch/), respectively. Harvestmen depicted in Figs. 1A–H,
consumed (Nyffeler et al. 2016). In recent years, however, new 2A, 3A–D, and 4A–B were identified by the following
evidence of vegetarian diet (i.e., plant and fungi consumption) in researchers: James C. Cokendolpher (Phalangiidae and
harvestmen and spiders has been published (e.g., Del-Claro et al. Sclerosomatidae), Adriano B. Kury (Cosmetidae, Gonylepti-
2017; Eastburn 2017; Lietzenmayer & Wagner 2017; Nahas et al. dae, Phalangiidae, and Sclerosomatidae), Ryosuke Kuwahara
2017; Hyodo et al. 2018; Benhadi-Marı́n et al. 2019; Cathrine (Sclerosomatidae), Glauco Machado (Cosmetidae and Gony-
2019; Pagoti et al. 2019; Suvák 2019; Hashimoto et al. 2020; leptidae), Jochen Martens (Gonyleptidae, Phalangiidae, and
Suzuki et al. 2021). The term ‘‘vegetarian’’ is currently used in Sclerosomatidae), Miguel Medrano (Cosmetidae), and Jeffrey
relation to carnivores, which occasionally switch to plant and/or W. Shultz (Sclerosomatidae). Spiders were identified by the
fungal food as an alternative to prey (e.g., Beckman & Hurd following researchers: Antonio D. Brescovit (Araneidae), G.B.
2003; Wäckers & Fadamiro 2005; Meehan et al. 2009). In this Edwards and David Hill (Salticidae), and Robert Raven
paper, we define ‘‘vegetarian food’’ as diets of plant or fungal (Sparassidae). Fungi depicted in Figs. 1A–H and 2C–D were
origin in addition to the arachnids’ consumption of prey. identified by the following researchers: Tim Baroni, Bart
Despite the increase in the records of fungal and plant Buyck, and Kathie T. Hodge (Russulaceae), Dennis E.
materials in the diet of harvestmen and spiders, many questions Desjardin and Serena Lee (Mycenaceae, Marasmiaceae, and
concerning their vegetarian diet remain unanswered. In Psathyrellaceae), Dennis E. Desjardin, Meike Piepenbring,
particular, it is still largely unexplored to what extent and Steven L. Stephenson (a Myxomycetes slime mold), and
harvestmen and spiders can be engaged in fungivory and Julieta Carranza (Pluteaceae). Fruits depicted in Figs. 3B, 3D,
frugivory. To fill this gap, we conducted a survey on this topic 3H, 4B–D, and Fig. S1 (online at https://doi.org/10.1636/JoA-
with the purpose of searching for reports of fungivory and S-22-015.s1) were identified by the following researchers:
frugivory in species of these two arachnid orders. In this paper, Maria do Carmo Amaral (Commelinaceae), Arik Hartmann
the term ‘‘frugivory’’ is used in a broad sense, by including the (Chrysobalanaceae), Samantha Koehler (Combretaceae),
consumption of fruit pulp, seeds, and seed appendages, such as Bruce Rottink (Fagaceae and Melanthiaceae), Renato Gold-
elaiosomes (i.e., fleshy structures, usually rich in lipids and enberg (Melastomataceae), Gilbert Barrantes (Piperaceae),
proteins, attached to the seeds of many plant species), and arils and Gustavo Shimizu (Primulaceae).
(i.e., a specialized outgrowth from a seed that partly or 2.3 Chemical property information for fungi and fruits.—The
completely covers the seed) (Rico-Gray & Oliveira 2008). Based following sources were used to obtain information on the
on the results of our review, we address four questions in our chemical properties of the fungal material and fruit species
discussion: (1) How common is fungivory and frugivory among consumed by harvestmen and spiders:
harvestmen and spiders? (2) Which taxa are engaged in
fungivory and frugivory? (3) What are the nutritional benefits – Todd & Bretherick (1942); Bartnicki-Garcia (1968); Fogel &
of fungivory and frugivory? (4) Can harvestmen and spiders be Trappe (1978); Barros et al. (2008); Chen et al. (2010) and
sustained on only fungal or fruit diets? Kalogeropoulos et al. (2013): Mushroom data for Canthar-
ellaceae, Marasmiaceae, Russulaceae, and other Basidio-
2. METHODS mycota; spore data on Basidiomycota / fungi Incerta sedis.
– United States Department of Agriculture database (online
2.1 Data collection.—We searched published reports on at https://fdc.nal.usda.gov/index.html): Fruit/Yeast data for
fungivory and frugivory in the Web of Science, Scopus, Adoxaceae, Bromeliaceae, Caricaceae, Cucurbitaceae, Fa-
Google Search, Google Scholar, Google Books, and Google gaceae, Juglandaceae, Musaceae, Rosaceae, Rutaceae, and
Pictures as well as ProQuest Dissertations and Theses, Saccharomycetaceae.
NYFFELER ET AL.—ARACHNIDS FEEDING ON FUNGI AND FRUITS 3
Table 1.—Reports of harvestmen and spiders feeding on mushrooms and other fungal material. In the column ‘‘Country’’, the letter (F)
indicates information gathered in the field and the letter (L) indicates information gathered in the laboratory. The column ‘‘Records’’ refers to the
number of written reports (e.g., papers, theses, and books) or internet images of a given arachnid species feeding on a fungus species. Records
followed by * were previously mentioned in the reviews of Acosta & Machado (2007) or Nyffeler et al. (2016).
No. of
Arachnid taxon Fungal species Fungal family Country records Reference
OPILIONES
Cladonychiidae
Erebomaster flavescens Non-identified Non-identified USA (F) 1* [1]
Cosmetidae
Non-identified Coprinellus sp. Psathyrellaceae Peru (F) 1 [2]
Globipedidae
Diguetinus raptator Non-identified Non-identified Mexico (F) 1 [3]
(decaying sporocarp)
Gonyleptidae
Iporangaia pustulosa Mycena sp. Mycenaceae Brazil (F) 1 [4]
Neosadocus sp. Non-identified Basidiomycetes Brazil (F) 1 [5]
Nemastomatidae
Nemastoma lugubre Non-identified Saccharomycetaceae England (L) 1* [6]
Nomoclastidae
Quindina albomarginis Non-identified Non-identified Myxomycetes Panama (F) 3 [7–9]
slime mold
Quindina limbata Pluteus sp. Pluteaceae Costa Rica (F) 1 [10]
Phalangiidae
Lophopilio palpinalis Cantharellus cibarius (sap) Cantharellaceae Czech Republic? (F) 1* [11]
Odiellus spinosus Non-identified Saccharomycetaceae England (L) 1 [6]
Oligolophus tridens Non-identified Saccharomycetaceae England (L) 1* [6]
Sclerosomatidae
Gagrella spinacantha Marasmius sp. Marasmiaceae Singapore (F) 1 [12]
Gagrella spinacantha Mycena sp. Mycenaceae Singapore (F) 1 [13]
Gagrella sp. Non-identified Mycenaceae Singapore (F) 1 [14]
Gagrellula ferruginea Mycena luxurius and Mycenaceae Japan (F) 1* þ 2 [15–17]
Mycena sp.
Leiobunum rotundum Non-identified Saccharomycetaceae England (L) 1 [6]
Leiobunum sp. Russula sp. Russulaceae USA (F) 2 [18–19]
Leiobunum spp. Non-identified Saccharomycetaceae USA (L) 1 [20]
Marthana niveata Marasmius sp. Marasmiaceae Malaysia (F) 1 [21]
Melanopa sp. Non-identified Non-identified Myxomicetes Japan (F) 1* [22]
slime mold
Pseudogagrella sp. Marasmius sp. Marasmiaceae Singapore (F) 1 [23]
Pseudogagrella sp. Mycena sp. Mycenaceae Malaysia (F) and Singapore (F) 5 [24–28]
Gagrellinae sp.1 Crinipellis sp. Marasmiaceae Singapore (F) 1 [29]
Gagrellinae sp.2 Hydropus sp. Mycenaceae Singapore (F) 1 [30]
Gagrellinae sp.3 Marasmius sp. Marasmiaceae Singapore (F) 1 [31]
Gagrellinae sp.4 Mycena sp. Mycenaceae Laos (F), Malaysia (F), and 4 [32–35]
Singapore (F)
Family not identified
Non-identified sp.1 Cantharellus sp. Cantharellaceae Scotland (F) 1 [36]
Non-identified sp.2 Mycena luxaeterna Mycenaceae Brazil (F) 1 [37]
Non-identified sp.3 Mycena sp. Mycenaceae Malaysia (F) 1 [38]
ARANEAE
Araneidae
Araneus diadematus Cladosporium herbarum Davidiellaceae Canada (F þ L) 2* [39–40]
Linyphiidae
Tenuiphantes tenuis Non-identified Saccharomycetaceae England (L) 1* [41]
[1] Goodnight & Goodnight (1960); [2] Photo by Rhett Butler, pers. comm. (Fig. 2A); [3] Cokendolpher et al. 2021; [4] John A. Uribe, pers.
comm. (see vı́deo: https://www.youtube.com/watch?v¼cjERQ2jSyRQ Accessed 16 June 2022); [5] Castanho & Rocha (2005); [6] Todd (1949); [7]
Rodrı́guez & Guerrero (1976); [8] Mora (1990); [9] Requena & Machado (2015), see also Fig. 2C; [10] Quesada-Hidalgo et al. (2019), see also Fig.
2D; [11] Šilhavý (1942); [12] Melvyn Yeo, pers. comm. (Fig. 1B); [13] Photo by Lenz Lim; [14] Melvyn Yeo, pers. comm.; [15] Uyemura (1935);
[16] Ryosuke Kuwahara, pers. comm. (Fig. 1F); [17] Anonymous photographer: http://fukuejima.la.coocan.jp/kinoko/shiino-tomoshibi-take.
html Accessed 19 April 2021; [18] Chuck Rosenberger, pers. comm. (Fig. 1H); [19] Anonymous photographer: https://www.shroomery.org/
forums/showflat.php/Number/26182555 Accessed 17 April 2021; [20] Edgar (1971); [21] Hock Ping Guek, pers. comm. (Fig. 1D); [22] Huzita
(1936); [23] Photo by Ang Kean Leng; [24] Hock Ping Guek, pers. comm. (Fig. 1E); [25] Anonymous photographer: https://www.gettyimages.ch/
detail/foto/macro-profile-shot-of-a-harvestman-feeding-on-lizenzfreies-bild/501778220?adppopup¼true Accessed 9 February 2022; [26–28]
Melvyn Yeo, pers. comm.; [29] Melvyn Yeo, pers. comm. (Fig. 1C); [30] Photo by Davin Tan; [31] Melvyn Yeo, pers. comm. (Fig. 1A); [32]
Photo by Marek Stefunko; [33] Melvyn Yeo, pers. comm. (Fig. 1G); [34] Photo by Thomas Calame; [35] Photo by Norjipin Saidi; [36] Bristowe
(1949); [37] Waldenmaier (2016); [38] Luping et al. (1978); [39] Smith (1984); [40] Smith & Mommsen (1984); [41] Sunderland et al. (1996).
Table 2.—Reports of harvestmen and spiders feeding on fruit pulp. In the column ‘‘Country’’, the letter (F) indicates information gathered in
the field and the letter (L) indicates information gathered in the laboratory. The column ‘‘Records’’ refers to the number of written reports (e.g.,
papers, theses, and books) or internet images of a given arachnid species feeding on the fruit pulp of a plant species. Records followed by * were
previously mentioned in the reviews of Acosta & Machado (2007) or Nyffeler et al. (2016).
Arachnid taxon Plant species Plant family Country No of records Reference
OPILIONES
Cosmetidae
Erginulus clavotibialis Ananas comosus Bromeliaceae Belize (L) 1 [1]
Eriginulus sp. Non-identified Non-identified Costa Rica (F) 1* [2]
Eupoecilaema magnum Piper sp. Piperaceae Costa Rica (F) 1 [3]
Gryne dimorpha Terminalia catappa Combretaceae Brazil (F) 1 [4]
Vonones ornatus Musa sp. Musaceae USA (L) 1 [5]
Globipedidae
Diguetinus raptator Opuntia sp. (rotten fruit) Cactaceae Mexico (F) 1 [6]
Diguetinus raptator Tuber of Pachyrhizus erosus or Fabaceae or Solanaceae Mexico (F) 1 [6]
Solanum tuberosum
Gonyleptidae
Acanthopachylus aculeatus Carica papaya Caricaceae Uruguay (L) 1 [7]
Discocyrtanus pertenuis Pyrus sp. (?) Rosaceae Brazil (L) 1 [8]
Discocyrtus invalidus Non-identified Non-identified Brazil (L) 1 [9]
Heteromitobates discolor Musa sp. Musaceae Brazil (L) 1 [10]
Iporangaia pustulosa Ardisia guianensis Primulaceae Brazil (F) 1 [11]
Iporangaia pustulosa Miconia blepharodes Melastomataceae Brazil (F) 1 [12]
Magnispina neptunus Terminalia catappa Combretaceae Brazil (F) 1 [13]
Mischonyx cuspidatus Musa sp. Musaceae Brazil (L) 2 [14–15]
Neosadocus bufo Eugenia strictosepala Myrtaceae Brazil (F) 1* [16]
Neosadocus bufo Non-identified Non-identified Brazil (F) 1* [17]
Opisthoplatus prospicuus Cucumis sativus Cucurbitaceae Uruguay (L) 1 [18]
Promitobates ornatus Carica papaya Caricaceae Brazil (L) 1 [19]
Serracutisoma proximum Miconia blepharodes Melastomataceae Brazil (F) 1 [20]
Nemastomatidae
Nemastoma lugubre Prunus domestica Rosaceae Denmark (L) 1 [21]
Oligophus tridens Prunus domestica Rosaceae Denmark (L) 1 [21]
Rilaena triangularis Prunus domestica Rosaceae Denmark (L) 1 [21]
Neopilionidae
Forsteropsalis bona Malus domestica Rosaceae New Zealand (L) 1 [22]
Forsteropsalis pureora Malus domestica Rosaceae New Zealand (L) 1 [22]
Forsteropsalis pureora Mangifera indica Anacardiaceae New Zealand (L) 1 [22]
Pantopsalis listeri Malus domestica Rosaceae New Zealand (L) 1 [22]
Phalangiidae
Phalangium opilio Quercus sp. (acorn) Fagaceae USA (F) 1 [23]
Sclerosomatidae
Gagrellula ferruginea Non-identified Non-identified Japan (F) 1 [24]
Leiobunum rotundum Rubus caesius Rosaceae England (F) 1 [25]
Leiobunum vittatum Rubus strigosus Rosaceae USA (F) 1* [26]
Leiobunum sp. Malus domestica Rosaceae USA (L) 1* [26]
Leiobunum sp. Malus domestica Rosaceae USA (L) 1 [27]
Leiobunum sp.1 Rubus fruticosus Rosaceae USA (F) 1* [28]
Leiobunum sp.2 Rubus sp. Rosaceae USA (F) 1 [29]
Leiobunum sp.3 Rubus sp. Rosaceae Netherlands (F) 1 [30]
Leiobunum sp.3 Sambucus nigra Adoxaceae Netherlands (F) 1 [30]
Non-identified sp. Rubus fruticosus Rosaceae USA (F) 1 [31]
ARANEAE
Amaurobiidae
Amaurobius similis Musa sp. Musaceae Scotland (F) 2 [32]
Araneidae
Alpaida leucogramma Carica papaya Caricaceae French Guinea (F) 1 [33]
Cheiracanthiidae
Cheiracanthium inclusum Chrysobalanus icaco Chrysobalanaceae USA (F) 1 [34]
Cheiracanthium sp. Citrus sinensis Rutatceae USA (F) 1 [35]
Philodromidae
Philodromus sp. Carica papaya Caricaceae Canada (F) 1 [36]
Table 2.—Continued.
Arachnid taxon Plant species Plant family Country No of records Reference
Salticidae
Phidippus apacheanus Citrullus lanatus Cucurbitaceae USA (L) 1 [37]
Phidippus audax Citrullus lanatus Cucurbitaceae USA (F) 1 [38]
Phidippus putnami Citrullus lanatus Cucurbitaceae USA (L) 1 [39]
Phidippus regius Citrullus lanatus Cucurbitaceae USA (L) 1 [40]
Unidentified web builders
Non-identified Rubus fruticosus Rosaceae England (L) 1 [41]
Non-identified Prunus domestica Rosaceae England (L) 1 [41]
[1] Schaus et al. (2013); [2] Acosta & Machado (2007); [3] Gilbert Barrantes, pers. comm. and Fig. 3B; [4] Mentioned as Paecilaema sp. in Pagoti
et al. (2019) and Fig. 3A; [5] Punzo et al. (2007); [6] Cokendolpher et al. (2021); [7] Capocasale & Bruno-Trezza (1964); [8] Photo by Joao Paulo
Burini and Fig. S1A; [9] dos Santos et al. (2013); [10] Costa et al. (2016); [11-12] Glauco Machado, pers. obs. and Figs. S1B –D; [13] Photo by
Rodrigo H. Willemart Fig. 3B; [14] Segovia et al. (2019); [15] Dı́as et al. (2020); [16] Machado & Pizo (2000); [17] Gnaspini (1996); [18] Fernandes
et al. (2017); [19] Willemart (2001); [20] Glauco Machado, pers. obs. and Figs. S1C–D; [21] Hvan & Toft (2008); [22] Powell et al. (2021); [23]
Photo by Darlene Watson https://www.pikist.com/free-photo-sbnqr Accessed 9 February 2022 and Fig. 3D; [24] Ryosuke Kuwahara, pers.
comm. and Fig. 3C; [25] Shardlow (2013); [26] Edgar (1971); [27] Schoffstall et al. (2013); [28] Halaj & Cady (2000); [29] Photo by Darren
Eugenius https://bugguide.net/node/view/1541187 Accessed 2 March 2022; [30] Wijnhoven (2011); [31] https://sarett.org/2021/07/10/nature-
notebook-daddy-longlegs/ Accessed 14 December 2021; [32] Cathrine (2019); [33] Sean McCann, pers. comm. and Fig. 1E; [34] Arik Hartmann &
Karl Nicolas Keiser, pers. obs. and Fig. 1H; [35] Isa Betancourt, pers. comm. and Fig. 1G; [36] Susannah Anderson, pers. comm. and Fig. 1F;
[37] https://www.reddit.com/r/spiders/comments/l8jknv/a_jumping_spider_eating_watermelon_they_can_gain/ Accessed 11 April 2021; [38]
https://www.facebook.com/photo/?fbid¼10104170091230947&set¼gm.5370362426315689 No longer available; accessed 19 May 2022; [39]
https://www.youtube.com/watch?v¼H2edfMCd3H8 Accessed 11 April 2021; [40] https://www.facebook.com/phantasticphids/videos/i-think-its-
adorable-how-excitedly-she-tapped-her-pedipalps-before-nomming-the-w/323934155482411/ Accessed 11 April 2021; [41] Bristowe (1941).
– Galetti et al. (2000): Fruit data for Euphorbiaceae, 3. RESULTS

Meliaceae, Myristicaceae, and Myrtaceae.
– Aguiar et al. (2011) and Maldonado et al. (2020): Fruit data In the following, we report on cases of fungivory and
for Chrysobalanaceae. frugivory in harvestmen and spiders. Consumption on seeds
– Udotong & Bassey (2015): Fruit data for Combretaceae. and seed appendages (elaiosomes and arils) was included in
– Messeder et al. (2021): Fruit data for Melastomataceae. the subsection on frugivory because seeds are a component of
– Foster (1977): Fruit data for Primulaceae. the fruits. A total of 96 records of fungivory/frugivory
– Herbst (1986) and Bizerril & Raw (1997): Fruit data for referring to harvestmen are reviewed in the present paper
Piperaceae. (Tables 1–3), 83% of which were not mentioned in the review
– Lisci et al. (1996) and Fischer et al. (2008): Elaiosome data of Acosta & Machado (2007). Twenty-eight records of
for Violaceae and other elaiosome-bearing plant taxa. fungivory/frugivory referring to spiders are presented (Tables
– Malacrida et al. (2011): Seed data for Caricaceae. 1–3), 86% of which were not found in the review of Nyffeler et
– Jordano (2000): Data for insects. al. (2016).
3.1 Fungivory.—In general, arachnids can consume mush-
2.4 Caloric value of fungi and fruits.—Caloric values for room tissue and sap, fungal spores, or yeast (Figs. 1–2).
various types of fungi and fruits were taken for the most part 3.1.1 Feeding on mushroom pileus or sap: Fungivory by
from the United States Department of Agriculture database. harvestmen has been known for a long time (e.g., Šilhavý
Additional data were extracted from Fogel & Trappe (1978), 1942; Bristowe 1949; Goodnight & Goodnight 1960). Con-
Herbst (1986), Bizerril & Raw (1997). Caloric values for sumption of fungal material has been witnessed on five
insects were adopted from Yhoung-Aree (2010). For some continents (Europe, North America, Central America, South
tropical fruits (i.e., Alchornea glandulosa, Ardisia sp., Cabralea America, and Asia). Most of the records (61%) refer to
canjerana, Eugenia stictosepala, Miconia sp., Terminalia tropical harvestmen in the family Sclerosomatidae (predom-
capatta, and Virola bicuhyba), which have no caloric values inantly subfamily Gagrellinae) (Table 1). The remaining cases
available in the literature, the values were estimated using the relate to the families Cladonychiidae, Cosmetidae, Gonylepti-
Atwater system (Mayxard 1944). This was done by adding up dae, Nomoclastidae, and Phalangiidae; in some cases, family
the calories provided by the energy-containing nutrients (i.e., identity is unknown (Table 1). Under field conditions, 64% of
fat, protein, and carbohydrate). The Atwater system is based the reports on fungivory included small forest mushrooms
on average values of 9 kcal/g for fat, 4 kcal/g for protein, and belonging to the families Marasmiaceae and Mycenaceae
4 kcal/g for carbohydrate (Food and Nutrition Information (dominated by Mycena spp., Figs. 1A–G), 17% of the reports
Center (FNIC), online at https://www.nal.usda.gov/legacy/ included mushrooms belonging to the families Cantharella-
fnic/how-many-calories-are-one-gram-fat-carbohydrate-or- ceae, Plutaceae, Psathyrellaceae (Fig. 2A), and Russulaceae
protein) and with knowledge of the fruits’ chemical compo- (Fig. 1H), and in 19% of the reports the identity of the family
sition (Table 4). Calculations were accomplished yielding was unknown (Table 1).
caloric values per 100 g of dry weight. Conversion to kcal/100 At least some of the fungi from the families Marasmiaceae
g of fresh weight was performed with knowledge of the fruits’ and Mycenaceae are bioluminescent in the dark (Fig. 2B).
water content (Table 4). Harvestmen are known to forage predominantly during the
Table 3.— Reports of harvestmen and spiders feeding on seeds (S) or seed appendages (E ¼ elaiosomes, A ¼ arils). In the column ‘‘Country’’,
the letter (F) indicates information gathered in the field and the letter (L) indicates information gathered in the laboratory (including
greenhouses). The column ‘‘Records’’ refers to the number of written reports (e.g., papers, theses, and books) or internet images of a given
arachnid species feeding on seeds or seed appendages of a plant species. Non-identified harvestmen sp.2†, sp.3†, and sp.8† were described by
Ohkawara & Higashi (1994) and Ohkawara et al. (1996, 1997) as spiders, but a picture sent to us by Ohkawara clearly shows that the animals in
question were harvestmen. Records followed by * were previously mentioned in the reviews of Acosta & Machado (2007) or Nyffeler et al.
(2016).
Arachnid taxon Plant species Plant family Seed type Country No. of records Reference
OPILIONES
Gonyleptidae
Neosadocus bufo Alchornea glandulosa Euphorbiaceae A Brazil (F) 1* [1]
Neosadocus bufo Cabralea canjerana Meliaceae A Brazil (F) 1* [1]
Neosadocus bufo Virola bicuhyba Myristicaceae A Brazil (F) 1* [1]
Serracutisoma proximum Dichorisandra paranaensis Commelinaceae A Brazil (F) 1 [2]
Phalangiidae
Oligolophus tridens Non-identified Non-identified S England (F) 1* [3]
(nut tree or shrub)
Sclerosomatidae
Leiobunum bimaculatum Erythronium americanum Liliaceae E USA (F) 1 [4]
Leiobunum sp.1 Carya sp. Juglandaceae S USA (F) 1* [5]
Nelima paessleri Trillium ovatum Melanthiaceae E USA (F) 1 [6]
Non-identified sp.1 Asarum canadense Aristolochiaceae E Canada (F) 1 [7]
Non-identified sp.2† Cordyalis ambigua Papaveraceae E Japan (F) 1 [8]
Non-identified sp.3† Erythronium japonicum Liliaceae E Japan (F) 1 [9]
Non-identified sp.4 Helianthus sp. Asteraceae S USA (F) 1 [10]
Non-identified sp.5 Jeffersonia diphylla Berberidaceae E USA (F) 1 [11]
Non-identified sp.6 Trillium spp. Melanthiaceae E USA (F) 1 [12]
Non-identified sp.7 Virola bicuhyba Myristicaceae A Brazil (F) 1 [13]
Non-identified sp.8† Viola spp. Violaceae E Japan (F) 1 [14]
ARANEAE
Araneidae
Neoscona adianta Non-identified Poaceae S France (F) 1* [15]
Zygiella sp. (?) Non-identified Non-identified S Norway (F) 1 [16]
Sparassidae
Heteropoda jugulans Carica papaya (seed) Caricaceae S Australia (F) 1 [17]
Theridiidae
Parasteatoda sp. Chelidonium majus Papaveraceae E Slovakia (L) 1 [18]
Parasteatoda sp. Costus dubius Costaceae E Slovakia (F) 1 [18]
Parasteatoda sp. Galanthus nivalis Amaryllidaceae E Slovakia (L) 1 [18]
Parasteatoda sp. Hepatica nobilis Ranunculaceae E Slovakia (L) 1 [18]
Uloboridae
Uloborus plumipes Asarum europaeum Aristolochiaceae E Slovakia (L) 1 [18]
Uloborus plumipes Chelidonium majus Papaveraceae E Slovakia (L) 1 [18]
Uloborus plumipes Cordyalis solida Papaveraceae E Slovakia (L) 1 [18]
Uloborus plumipes Costus dubius Costaceae E Slovakia (L) 1 [18]
Uloborus plumipes Galanthus nivalis Amaryllidaceae E Slovakia (L) 1 [18]
Uloborus plumipes Hepatica nobilis Ranunculaceae E Slovakia (L) 1 [18]
[1] Machado & Pizo (2000); [2] Bruno A Buzatto & G. Machado, pers. obs.; [3] Bristowe (1949); [4] Ruhren & Dudash (1996); [5] Wickham
(1918); [6] Bruce Rottink, pers. comm.; [7] Dunphy et al. (2016); [8] Ohkawara et al. (1997); [9] Ohkawara et al. (1996); [10] Photo by M.J.
Hatfield: https://bugguide.net/node/view/1940496 Accessed 17 April 2021; [11] Smith et al. (1986); [12] Gunther & Lanza (1989); [13] Zipparro &
Morellato (2005); [14] Ohkawara & Higashi (1994); [15] Berland (1933); [16] Olav Berge Aamodt, pers. comm.; [17] https://www.facebook.com/
YahooNewsAu/videos/spider-drinking-from-papaya-seed/727544364558928/ [posted 3 December 2020] Accessed 12 March 2022; [18] Suvák
(2019).
night hours (Acosta & Machado 2007) and appear to be bioluminescent glow-worms (Meyer-Rochow & Liddle 1988,
attracted by the light emission of bioluminescent mushrooms 2001).
(Fig. 2B; Luping et al. 1978; Waldenmaier 2016). In fact, it has There are reports of fungivory among parental males of two
been experimentally demonstrated that the New Zealand species of the genus Quindina Roewer, 1915 (Nomoclastidae)
harvestmen Forsteropsalis tumida (Forster, 1944) (Neopilioni- (Rodrı́guez & Guerrero 1976; Mora 1990; Requena &
dae) and Hendea myersi (Phillipps & Grimmett, 1932) Machado 2015; Rojas et al. 2019). In this genus, males build
(Triaenonychidae) are attracted to the light emitted by mud nests on fallen tree trunks that are used by females as
Figure 1.—Harvestmen feeding on mushrooms. A. Unidentified Gagrellinae (Sclerosomatidae) feeding on the mushroom Marasmius sp.
(Marasmiaceae) in Singapore (Photo by Melvin Yeo). B. Gagrella spinacantha Roewer, 1954 (Sclerosomatidae, Gagrellinae) feeding on the
mushroom Marasmius sp. (Marasmiaceae) in Singapore (Photo by Melvin Yeo). C. Unidentified Gagrellinae (Sclerosomatidae) feeding on the
mushroom Crinipellis sp. (Marasmiaceae) in Singapore (Photo by Melvin Yeo). D. Marthana niveata (Roewer, 1955) (Sclerosomatidae,
Gagrellinae) feeding on the mushroom Marasmius sp. (Marasmiaceae) in Malaysia (Photo by Hock Ping Guek). E. Pseudogagrella sp.
(Sclerosomatidae, Gagrellinae) feeding on a mushroom (Mycenaceae) in Malaysia (Photo by Hock Ping Guek). F. Gagrellula ferruginea (Loman,
1902) (Sclerosomatidae, Gagrellinae) in the process of tearing off the pileus (’0.5 cm in diameter) of the mushroom Mycena sp. (Mycenaceae) in
Japan (Photo by Ryosuke Kuwahara). G. Unidentified Gagrellinae (Sclerosomatidae) feeding on the mushroom Mycena sp. (Mycenaceae) in
Singapore (Photo by Melvin Yeo). H. Leiobunum sp. (Sclerosomatidae, Leiobuninae) feeding on the mushroom Russula sp. (Russulaceae) in
Hart County, Kentucky, USA (Photo by Chuck Rosenberger).
Figure 2.—Further cases of harvestmen feeding on mushrooms. A. Unidentified Cosmetidae feeding on the mushroom Coprinellus sp.
(Psathyrellaceae) in Tambopata area, Peru (Photo by Rhett Butler). B. Bioluminescent mushrooms growing on the forest floor in Singapore emit
a greenish light at night. Unidentified harvestmen were often seen feeding on the pileus of such bioluminescent mushrooms (Photo by Nicky
Bay). C. Male of Quindina albomarginis (Chamberlin, 1925) (Nomoclastidae) inside his mud nest built on a fallen log in Panama (the dorsum and
hind legs are marked with colored ink). Note that, by feeding on the growing fungi (a Myxomycetes slime mold), the parental male keeps the nest
floor and the eggs cleaner when compared to the areas outside the nest (Photo by Gustavo S. Requena). D. Male of Quindina limbata (Roewer,
1943) (Nomoclastidae) inside his mud nest built on a fallen log in Costa Rica. Again, the parental male keeps the nest and the eggs clean by
feeding on the fungi that grow on the fallen trunk (Photo by Andrés Rojas).
oviposition sites. The nests are usually infested by fungi, which Mommsen 1984). The digestive fluid of spiders contains the
can kill the eggs (Mora 1990; Quesada-Hidalgo et al. 2019; enzyme chitinase needed to dissolve and digest the chitinous
Rojas et al. 2019). Males feed on the fungi, keeping the nest spore cell wall, which explains why spiders can digest fungal
clean and the eggs alive (Figs. 2C–D). spores (Mommsen 1978, 1980; Smith & Mommsen 1984;
Spiders are frequently found resting near or on mushrooms Nyffeler et al. 2016). Spores from many different fungal
(see the large number of examples uploaded on Google families (e.g., Botryosphaeriaceae, Davidiellaceae, Helotia-
Pictures using the key words ‘‘spider þ mushroom’’), but they ceae, Massarinaceae, Microascaceae, Nectriaceae, Phragmi-
have never been reported feeding on the pileus or stipe of diaceae, Pleosporaceae, Trichocomaceae, Trichosphaeriaceae,
mushrooms. This was confirmed by a nocturnal infrared video and Venturiaceae) are blown by wind into spider webs (Smith
surveillance study conducted in the Atlantic Rainforest near & Mommsen 1984; Linskens et al. 1993; Bera et al. 2002; Del
São Paulo, southeastern Brazil, in two consecutive years Fiol et al. 2007; Quamar & Chauhan 2011; Nyffeler et al.
(Waldenmaier 2016). According to this surveillance study, 2016). The spores trapped in spider webs belong to the most
ground spiders from the families Gnaphosidae and Lycosidae
common fungal genera one would expect to find in the air
frequently visited the area of Mycena luxaeterna, staying there
(Nyffeler et al. 2016). In Italy, Del Fiol et al. (2007) found
for up to 30 min, but always without feeding on the
’17,000 fungal spores trapped in six orb-webs of the spider A.
mushrooms (Waldenmaier 2016). The only mentions of
diadematus (sampled from summer of one year to spring of the
naturally occurring fungivory in spiders are cases of ingestion
of fungal spores by orb-weavers (see next subsection). next year), leading to the conclusion that fungal spores might
3.1.2 Feeding on fungal spores and yeast: Currently, reports be a supplementary non-prey food of some nutritional
of consumption of fungal spores by harvestmen are lacking importance (but see subsection ‘‘What are the nutritional
(see Acosta & Machado 2007; Lundgren 2009). Spiders, on the benefits of fungivory and frugivory?’’ in Discussion). Besides
other hand, get access to fungal spores as a food source if fungal spore consumption, linyphiid spiders consumed yeast
airborne fungal spores blown by wind into their webs get stuck material in laboratory feeding trials (Sunderland et al. 1996).
to the viscid threads (Smith & Mommsen 1984; Linskens et al. Consumption of yeast under laboratory conditions was
1993; Del Fiol et al. 2007). The spores are ingested along with observed in harvestmen as well (Todd 1949; Edgar 1971).
the old web prior to the construction of a new web, as has been 3.2 Frugivory.—This type of feeding behavior refers to the
shown in laboratory experiments with the orb-weaver Araneus consumption of fruit pulp, seeds, and seed appendages, such
diadematus Clerck, 1757 (Araneidae) (Smith 1984; Smith & as elaiosomes and arils (Figs. 3–4). While frugivory by
Figure 3.—Harvestmen and spiders feeding on fruits. A. The harvestman Eupoecilaema magnum Roewer, 1933 (Cosmetidae) feeding on a ripe
Piper sp. fruit directly on the shrub in Costa Rica (Photo by Gilbert Barrantes). B. Three individuals of the harvestman Gryne dimorpha Mello-
Leitão 1928 (Cosmetidae) and one individual of the harvestman Magnispina neptunus Mendes, 2011 (Gonyleptidae) feeding on a fallen fruit of
Terminalia catappa (Combretaceae) in Northeastern Brazil (Photo by Rodrigo H. Willemart). C. The harvestman Gagrellula ferruginea (Loman,
1902) (Sclerosomatidae) feeding on an unidentified wild fruit in Japan (Photo by Ryosuke Kuwahara). D. The harvestman Phalangium opilio
Linnaeus, 1761 (Phalangiidae) feeding on an oak acorn fruit of Quercus sp. (Fagaceae) in USA (Photo by Darlene Watson). E. The orb-weaving
spider Alpaida leucogramma (White, 1841) (Araneidae) feeding on a piece of a papaya fruit waste (Caricaceae) thrown into a yard in Cayenne,
French Guiana (Photo by Sean McCann). F. The spider Philodromus sp. (Philodromidae) feeding on a speck of papaya pulp left on the kitchen
counter in a residence in Campbell River, British Columbia, Canada (Photo by Susannah Anderson). G. The yellow sac spider Cheiracanthium
sp. (Cheiracanthiida) feeding on a slice of blood orange (Rutaceae) that had been left on a kitchen counter cutting board in Ann Arbor,
Michigan, USA (Photo by Isa S. Betancourt). H. The yellow sac spider Cheiracanthium inclusum (Cheiracanthiidae) feeding on an overripe
cocoplum fruit of Chrysobalanus icaco (Chrysobalanaceae) in the Everglades, USA (Photo by Arik Hartmann).
Figure 4.—Seeds bearing elaiosomes and arils that are consumed by predatory arthropods, including harvestmen and spiders. A. Harvestman
– presumably an immature of Nelima paessleri (Roewer, 1910) (Sclerosomatidae) – feeding on elaiosomes of Trillium ovatum (Melanthiaceae) in a
forested area in Oregon, USA (Photo by Bruce Rottink). B. Open capsule of the fruit of Dichorisandra paranaensis (Commelinaceae) showing the
seeds surrounded by a white aril. The aril of one of these seeds is chewed by the harvestman Serracutisoma proximum (Mello-Leitão 1922)
(Gonyleptidae) in southeastern Brazil (Photo by Bruno A. Buzatto). C. Seed of Cabralea canjerana (Meliaceae) fallen on the leaf litter in the
Brazilian Atlantic forest. Seeds of this species are surrounded by a lipid-rich aril that is consumed by several ant species (such as the Pheidole sp.
depicted in the photo) and by the harvestman Neosadocus bufo (Photo by Marco Aurélio Pizo). D. Seed of Virola bicuhyba (Myristicaceae) fallen
on the leaf litter in the Brazilian Atlantic forest. Seeds of this species are also surrounded by lipid-rich aril that is consumed by ants (such as the
Pheidole sp. depicted in the photo) and by the harvestman N. bufo (Photo by Marco Aurélio Pizo). E. The spider Parasteatoda sp. (Theridiidae)
feeding on elaiosome-bearing seed of Costus dubius (Costaceae) in a greenhouse in Slovakia (Photo by Martin Suvák). F. The spider Parasteatoda
sp. (Theridiidae) feeding on elaiosome-bearing seed of Hepatica nobilis (Ranunculaceae) during a greenhouse experiment (Photo by Martin
Suvák).
harvestmen has been known for a long time, frugivory by Gonyleptidae fed on the pulp of fallen fruits (Gnaspini 1996;
spiders is a new area of research (see also Nyffeler et al. 2016). Machado & Pizo 2000; Acosta & Machado 2007; Pagoti et al.
3.2.1 Feeding on fruit pulp: Consumption of fruit pulp by 2019; Fig. 3B; Fig. S1C). In two cases, species from the
harvestmen has been reported from six continents (Europe, families Cosmetidae and Gonyleptidae were also observed
Asia, Oceania, North America, Central America, and South feeding on ripe fruits directly on trees and shrubs (Fig. 3A;
America), and was observed in seven families (Cosmetidae, Fig. S1B).
Globipedidae, Gonyleptidae, Nemastomatidae, Neopilioni- In the laboratory, there are also records of harvestmen
dae, Phalangiidae, and Sclerosomatidae; Table 2). In the consuming fruit pulp (Table 2). In the Holarctic region,
Holarctic region, harvestmen in the genus Leiobunum (Scle- Lacinius dentiger (Koch, 1847) (Phalangiidae) accepted apples
rosomatidae) were repeatedly seen feeding on Rubus spp. and pears (Mitov 1988), and several species of neopilionids
(Rosaceae) berries (Edgar 1971; Halaj & Cady 2000; also accepted apples (Table 2). In the Neotropical region, the
Wijnhofen 2011; Shardlow 2013; see also Bugguide, online gonyleptids Heteromitobates discolor (Sørensen, 1884) and
at https://bugguide.net/node/view/1541187). In Neotropical Mischonyx squalidus Bertkau, 1880 accepted banana (Costa et
forests, harvestmen from the families Cosmetidae and al. 2016; Segovia et al. 2019; Dias et al. 2020), Opisthoplatus
Table 4.—Major chemical properties and caloric value (kcal/100 g) for various taxa of fungi and fruits used as food by arachnids (see Tables
1–3). Abbreviations: A ¼ Araneae (spiders); O ¼ Opiliones (harvestmen); E (superscript) ¼ information obtained in experiments; FW ¼ fresh
weight; DW ¼ dry weight; # ¼ combined value for proteins þ amino acids; * ¼ values roughly estimated using the Atwater system (see Methods
subsection 2.4); ** ¼ chemical composition of the fleshy aril; NA ¼ non-available information.
Arachnid Water Lipids Protein Carbohydrates kcal/100 Avg. fruit weight

Fungus/Plant family order Fungus or fruit species % FW % DW % DW % DW g FW (g FW/fruit)
FUNGI
Cantharellaceae O Cantharellus sp. 92 2 54 32 28 –
Marasmiaceae O Marasmius sp. ~90 3 40 34 ~38 –
Russulaceae O Russula sp. 90 5–7 20–30 50–60 39 –
Basidiomycota O Various spp. 70–94 1–7 12–62 28–70 30–50 –
Basidiomycota A Unspecified spores 15 1–10 10–16 30–60 335* –
Saccharomycetaceae A Unspecified species 70 8 42 43 116* –
FRUITS (PULP)
Adoxaceae O Sambucus nigra 80 2.5 3 91 73 ,1
Bromeliaceae OE Ananas comosus 86 1 4 94 50 1,600
Caricaceae A, OE Carica papaya 88 2 4 91 43 450
Chrysobalanaceae A Chrysobalanus icaco 79–84 3.5 3 91 84 9.5
Combretaceae O Terminalia catappa 81 1 9 83 70 25
Cucurbitaceae OE Cucumis sativus 95 2 13.5 76 15 400
Cucurbitaceae AE Citrullus lanatus 91 2 7 88 30 3,500
Fagaceae O Quercus sp. (acorn) 28 33 8.5 57 236 3
Melastomataceae O Miconia sp. 81 7.5 12 68 74* ,1
Musaceae A, OE Musa sp. 75 1.5 4 91 89 120
Myrtaceae O Eugenia stictosepala 78 5 8.5 85.5 ~90* 6
Piperaceae O Piper spp. 73–78 1.5–14 6–9 52–87 35–107 1–9
Primulaceae O Ardisia sp. (mesocarp) 88 3 2 82 44* ,1
Rosaceae OE Malus domestica 86 1 2 96 52 70–100
Rosaceae AE, OE Prunus domestica 87 2 5.5 89 46 65
Rosaceae OE Pyrus spp. 84 1 2 95 57 200
Rosaceae O Rubus caesius 88.5 4 12 83.5 43 4
Rosaceae O Rubus fruticosus 88 4 12 81 43 2.5
Rosaceae O Rubus idaeus/strigosus 86 4.5 8.5 84 52 5
Rutaceae A Citrus sinensis 87 1 7 89 45 130
SEEDS AND
SEED APPENDAGES
Caricaceae A Carica papaya 44 29 26 30.5 NA –
Euphorbiaceae O Alchornea glandulosa 43** 68** 8** 22** ~400* ,1
Juglandaceae O Carya sp. (nut) 3 66 13 19 190 3–12
Meliaceae O Cabralea canjerana 48** 71** 10** 16.5** ~400* 1
Myristicaceae O Virola bicuhyba 63** 62** 5** 32** ~250* 3.5
#
Various families A, O Various species bearing 86 ~60 ~28 ~11 160–230 –
elaiosomes (mean value)
Violaceae O Viola spp. elaiosomes NA NA NA NA 170–190 –
INSECT PREY
Insecta – Various taxa (mean value) 64 17 68 15 98–231 –
prospicuus (Holmberg, 1876) accepted cucumber (Fernandes et or orange pulp) left over on kitchen counters or disposed in
al. 2017), Discocyrtanus pertenuis (Mello-Leitão, 1935) accept- yards (Table 2; Figs. 3E–G).
ed pear (Fig. S1A), and Acanthopachylus aculeatus (Kirby, Under field conditions, an American yellow sac spider,
1819) and Promitobates ornatus (Mello-Leitão, 1922) accepted Cheiracanthium inclusum Hentz, 1847 (Cheiracanthiidae), was
papaya (Capocasale & Bruno-Trezza 1964; Willemart 2001). observed feeding on an overripe cocoplum fruit (Chrysobala-
More recently, several cases of free-living spiders feeding on nus icaco; Chrysobalanaceae) in the Everglades National Park,
fruits have been witnessed (Table 2). In central Scotland, Florida, USA (Fig. 3H). Chrysobalanus icaco is a common
Cathrine (2019) witnessed on several occasions males of native shrub in the Everglades ecosystem. Continued obser-
Amaurobius similis (Blackwall, 1861) (Amaurobiidae) feeding vation showed the spider manipulating the pulp of the fruit
on remains of banana fruit pulp (Musa sp.; Musaceae) left on with its chelicerae (see https://figshare.com/articles/media/
a kitchen table. This unusual feeding behavior in Amaurobius American_yellow_sac_spider_Cheiracanthium_inclusum_
was witnessed over several years in two residencies located Cheiracanthiidae_feeding_on_an_overripe_cocoplum_fruit_
about 11 km apart (Cathrine 2019). There are similar reports Chrysobalanus_icaco_Chrysobalanaceae_/19349174 Accessed
from North America and South America of free-living spiders March 10th 2022). It is unknown if the spider employed
feeding on fruit remains (e.g., pieces of papaya, watermelon, digestive enzymes to liquefy the flesh of the fruit, or if the
ripened flesh was of appropriate consistency to be imbibed by cally feed on small pieces of fruit waste encountered inside
the spider. houses or disposed in yards (Tables 2–3). Consumption of seed
3.2.2 Feeding on seeds and seed appendages: We found 16 appendages, in turn, has been reported in a greenhouse
records of harvestmen feeding on seeds or seed appendages experiment with two spider species (Table 3), and it is not
(Table 3). These reports refer in most cases (’81%) to the possible to know how frequent this feeding habit is under
consumption of elaiosomes and arils. In the Holarctic region, natural field conditions.
harvestmen were observed consuming the elaiosomes of plants 4.2 Which taxa are engaged in fungivory and frugivory?—In
in the families Aristolochiaceae, Berberidaceae, Liliaceae, the survey by Acosta & Machado (2007), information on the
Melanthiaceae, Papaveraceae, and Violaceae (Table 3; Fig. diet of harvestmen is available for 13 families (’24% of all
4A). In the Neotropical region, there is a record of the living families) and vegetarian feeding habits (ie., consumption
gonyleptid Serracutisoma proximum (Mello-Leitão, 1922) of fungi, fruits, seeds, and seed appendages) have been
feeding on the aril of a Commelinaceae (Fig. 4B). Moreover, documented for four families: Cladonychiidae, Gonyleptidae,
the gonyleptid Neosadocus bufo (Mello-Leitão, 1923) has been Phalangiidae, and Sclerosomatidae. In the present review, we
regularly observed feeding on the fleshy arils of the capsular expand the number of harvestman families with records of
fruits of Alchornea glandulosa, Cabralea canjerana, and Virola vegetarian feeding, which now also includes Cosmetidae,
bicuhyba (Machado & Pizo 2000). The arils of these three Globipedidae, Nemastomatidae, Neopilionidae, and Nomo-
plant species become exposed to ground dwelling arthropods, clastidae (Tables 1–3). Most of the documented feeding events
such as ants and harvestmen, after the capsules open and the summarized here refer to the Sclerosomatidae and Gonylepti-
seeds fall on the forest floor (Figs. 4C–D). dae, which are the largest harvestman families (Machado et al.
Several records of spiders seen feeding on seeds or seed 2007; Kury et al. 2021). Considering that 54 living families
appendages were reported in recent years (Table 3). Berland exist in the order Opiliones, it follows that little is known
(1933) was perhaps the first to report an araneid orb-weaving whether species of 45 families also engage in vegetarian
spider, Neoscona adianta (Walckenaer, 1802), sucking a grass feeding. There are two non-mutually exclusive explanations
seed (Poaceae). A similar incident – also concerning an araneid for this gap in our knowledge. First, some harvestman taxa
orb-weaving spider feeding on a plant seed – was witnessed apparently are exclusively carnivorous feeders (e.g., families
recently in a residential area of the city of Sandnes, Norway Ischyropsalididae and Trogulidae; see Nyffeler & Symondson
(Olav Berge Aamodt, pers. comm.). Moreover, a huntsman 2001 and Acosta & Machado 2007). Second, the feeding habits
spider, Heteropoda jugulans (L. Koch, 1876) (Sparassidae), of most harvestman taxa are still unexplored (Powell et al.
was filmed in the kitchen of a residence in Brisbane, 2021). However, based on anecdotal evidence accumulated so
Queensland, Australia, feeding on a papaya seed that had far, we anticipate that consumption of fungi, fruit pulp, seeds,
been removed from a cutting board (Yahoo News Australia
and seed appendages is widespread in many harvestman
2020). Finally, in a greenhouse in Slovakia, the comb-footed
families, especially those that inhabit tropical forests, where
spider (Parasteatoda sp., Theridiidae) was seen feeding on
the availability of these food items is probably high over most
elaiosomes of the elaiosome-bearing seeds of Costus dubius
parts of the year, including the dry and cold season, when
(Costaceae) trapped in its web (Fig. 4E; Suvák 2019). To learn
arthropod prey are scarcer (Wolda 1988).
more about the capability of spiders to use elaiosomes as a
For spiders, the situation is somewhat different. So far,
potential food source, Suvák (2019) conducted a greenhouse
fungivory has been documented under natural conditions
experiment by throwing elaiosome-bearing seeds of various
exclusively for ecribellate orb-weaving spiders (family Aranei-
plant taxa into webs of the spiders Parasteatoda sp. and
dae), which digest airborne fungal spores during the recycling
Uloborus plumipes Lucas, 1846 (Uloboridae). This experiment
revealed that both species fed readily on the elaiosomes (Table process of old webs prior to the construction of new webs
3; Fig. 4F). (Smith & Mommsen 1984; Table 1). It is possible that web-
building spiders other than orb-weavers also occasionally feed
on fungal material trapped in their webs, as the laboratory
4. DISCUSSION
feeding trials of Sunderland et al. (1996) suggest. Frugivory, in
4.1 How common is fungivory and frugivory in harvestmen turn, seems to occur predominantly among synanthropic
and spiders?—Consumption of fungal material by harvestmen spiders from different families that opportunistically encoun-
has been reported more frequently in warmer areas, with ter small pieces of fruit waste mostly inside houses (Table 2).
.70% of all reports originating from tropical locations, Frugivory by spiders under natural field conditions is
especially Southeast Asia (Table 1). In contrast to harvestmen, documented here for the first time, including one species of
consumption of fungal material by spiders seems to be less the genus Cheiracanthium (Table 2). This result could be
frequent (Table 1). However, consumption of fungal spores by biased by the synanthropic spiders being probably more often
spiders does occur all over the world and is not limited to a observed by humans, when compared with the wild ones.
specific region (e.g., Smith 1984; Smith & Mommsen 1984; Finally, the few cases of spiders feeding on seed appendages
Bera et al. 2002; Del Fiol et al. 2007; Walter et al. 2019). reported here (Table 3) appear to be uncommon occurrences,
Feeding on fruit pulp, seeds, and seed appendages (i.e., restricted to greenhouse conditions. In the last three decades, a
elaiosomes and arils) by harvestmen was reported from areas large number of quantitative prey analyses on spiders have
of temperate and tropical climates (Tables 2–3). Feeding on been conducted without detecting them feeding on fruit pulp,
fruit pulp and seeds by spiders was also reported from seeds, or seed appendages (see Nentwig 1987; Wise 1993;
different parts of the world, but this appears to be largely Nyffeler 1999; Pekár & Toft 2015). Thus, we suggest that,
limited to cases in which synanthropic species opportunisti- although frugivory by spiders may have been overlooked by
arachnologists in the past, this type of feeding habit is harvestmen and spiders have been reported to feed on fruit
probably rare in this order. pulp refer to low-quality fruits (Table 4). This pattern raises
4.3 What are the nutritional benefits of fungivory and the question whether feeding on fruits of low caloric value is of
frugivory?—As pointed out in the Results, harvestmen and any nutritional benefit. Cathrine (2019), who observed spiders
spiders use fungal material in different ways. Harvestmen that feeding on banana waste, commented ‘‘Whether this is for
feed on fungal tissues (especially mushrooms) are provided sustenance or moisture is unclear. . .’’. Several researchers
with an abundant supply of water, carbohydrates, protein, conducted food choice experiments under laboratory condi-
minerals, and vitamins (Lundgren 2009). In particular, the tions and these experiments may provide an answer to the
high protein content (usually ’20–40% on a dry weight basis; question raised above.
Table 4) of mushroom tissues is remarkable, but compared to Schaus et al. (2013) carried out a feeding trial in which the
insect prey, mushroom tissues have a low caloric value (’30– Neotropical harvestman Erginulus clavotibialis (Pickard-Cam-
50 kcal/100 g of fresh weight; Table 4). Furthermore, some bridge, 1905) (Cosmetidae) was given a choice between fresh
mushrooms (e.g., Russula spp.) contain toxic substances pineapple (i.e., low-quality fruit) and live invertebrate prey.
(Matsuura et al. 2016), but nothing is known of harmful This harvestman demonstrated a distinct preference for fruit
effects on harvestmen caused by the consumption of toxic over invertebrate prey, clearly suggesting that feeding on fresh
mushrooms. fruit was nutritionally beneficial to the individuals even
The situation is very different regarding the consumption of though the offered fruit was of the low-quality type.
fungal spores by spiders. Spores have a high caloric value Furthermore, Schaus et al. (2013) showed that individuals
(’300 kcal/100 g of fresh weight; Table 4) accompanied by a feeding on fruits were more active compared to starved
low water content (’15%). Contrary to fungal tissues, the individuals or individuals that fed on live invertebrate prey.
protein content of the spores is rather low (’10–16% on a dry Analogous food choice experiments were conducted with
weight basis; Table 4), which renders them as food of very low spiders in the families Salticidae, Thomisidae, and Any-
or no nutritive value to spiders (Smith & Mommsen 1984; phaenidae (Vogelei & Greissl 1989; Pollard et al. 1995;
Lundgren 2009; Parish et al. 2020). Smith & Mommsen (1984) Jackson et al. 2001; Taylor & Bradley 2009; Pfannenstiel &
have shown in laboratory feeding trials that second instar Patt 2012). In these experiments, the spiders were permitted to
spiderlings of the orb-weaving spider Araneus diadematus choose between a sucrose solution and distilled water. In all
which had access to fungal spores as a potential food source experiments the spiders spent more time drinking from the
did not differ significantly in their chance of survival from a sucrose solution compared to water, showing that the spiders
starved control group. fed on sugar solutions to obtain nutrients and not just
The suitability of fungal spores as a diet might be further moisture. In these experiments, it has also been shown that
compromised by the presence of noxious secondary com- consumption of sucrose solution increased spider survival
pounds (Smith & Mommsen 1984; Lundgren 2009). An significantly compared to consumption of water only. If we
experiment in which a mixed diet of plant pollen and yeast extrapolate these experimental findings to the situation of
(family Saccharomycetaceae) was offered to early instar harvestmen and spiders feeding on low-quality fruits under
spiderlings of the sheet-web spider Tenuiphantes tenuis (Black- natural field conditions, we conclude that they feed on fruits
wall, 1852) (Linyphiidae) confirmed the hypothesis that the probably to obtain nutrients in addition to moisture.
ingestion of fungi has a detrimental effect. Spiderlings feeding Not only do harvestmen and spiders feed on fruit pulp, but
on the pollen-yeast diet showed a 50% decrease in survival they also feed on the fruits’ most inner parts, the seeds and,
time compared to a group of starved spiderlings (Sunderland more frequently, their appendages. In fact, most cases in
et al. 1996). As a final remark, we stress that it is currently which harvestmen and spiders were documented feeding on
unknown whether the consumption of fungal spores plays a seeds refer to the consumption of elaiosomes or arils (Table 3).
role in the dynamics of araneopathogenic fungi whose spores The nutritional quality of elaiosomes and arils as food
are infective propagules (Durkin et al. 2021). This would resembles that of insects as regards their fatty acid composi-
require spores to come into contact with, and adhere to, the tion (Hughes et al. 1994). Moreover, the lipid content of
host cuticle during consumption, as infection from inside the elaiosomes and arils is usually very high (Rico-Gray &
gut is unlikely (as far as we know). Oliveira 2008; see also Table 4). For instance, the arils of
Fruits can be classified into high-quality and low-quality Alchornea glandulosa, Cabralea canjerana, and Virola bicuhyba
according to their caloric value (Johnson et al. 1985). High- contain a much higher percentage of lipids (62–71%) when
quality fruits usually have a water content of ’40–60%, are compared to the average value of various insect prey, which is
rich in lipids (’60–70% on a dry weight basis) and have a high around 17% (see Table 4). Various insect groups, particularly
caloric value (’250–400 kcal/100 g of fresh weight; Table 4). ants, are known to feed on elaiosomes and arils (Lundgren
In turn, fruits considered to be of low quality, are less 2009; see also Figs. 4C–D). Here we showed that elaiosomes
nutritious (the pulp containing , 10% lipid on a dry weight and arils are also consumed by arachnids. Whereas consump-
basis; Table 4) with a watery flesh (’70–90% moisture; Table tion of elaiosomes and arils by harvestmen is probably
4). Such fruits have a high content of carbohydrates, some of widespread, it has been reported for only a few spider genera
which are in the form of sugars (i.e., fructose, glucose, and in two families, always under artificial conditions (Table 3).
sucrose). Fruits of this type (e.g., apple, banana, berries, 4.4 Can harvestmen and spiders be sustained on only fungal or
cocoplum, cucumber, elderberry, orange, papaya, pear, plum, fruit diets?—On Fukue Island and in other regions of Japan,
pineapple, and watermelon) are a poor source of energy (’15– some harvestman species (eg., Gagrellula ferruginea) are often
70 kcal/100 g of fresh weight; Table 4). Most cases in which seen feeding on the pileus of luminescent mushrooms (Mycena
luxurius and Mycena spp.) growing on the forest floor (Fig. whenever fruits, seeds, and seed appendages are abundant
1F; Uyemura 1935; http://fukuejima.la.coocan.jp/kinoko/ (Machado & Pizo 2000; Schaus et al. 2013; Pagoti et al. 2019).
shiino-tomoshibi-take.html; Ryosuke Kuwahara, pers. Harvestmen are considered to be diet generalists and feeding
comm.). Likewise, harvestmen were reported eating on on mixed diets of plants and invertebrate prey is in several
luminescent mushrooms of the family Mycenaceae growing ways advantageous because: (1) it broadens the trophic niche,
on the ground of lowland rain forests in Malaysia (Luping et (2) it attenuates competition for food with other predatory
al. 1978) and Singapore (Melvin Yeo, pers. comm.). Further- arthropods, (3) it allows to switch increasingly to plant
more, in the Brazilian Atlantic Forest, unidentified harvest- resources at times when invertebrate prey becomes scarce, and
men were observed feeding on pileus tissue of the mushroom (4) it dilutes adverse effects of toxic compounds contained in
Mycena luxaeterna, which emits intense light during the night some food sources (see Coll & Guershon 2002). However,
(Waldenmaier 2016). In all these examples, harvestmen were when harvestmen were fed an only plum diet (Prunus
reported to feed heavily on mushrooms, indicating that domestica), they lost weight compared to harvestmen fed a
fungivory may play an important role in the diet of certain diet containing arthropod prey (Hvam & Toft 2008). This
species. indicates that, although harvestmen are rather generalist
The genus Quindina is an interesting model system to study feeders, they may be unable to survive on a diet composed
fungivory in harvestmen. During the period of paternal care, exclusively of low-quality fruits, such as plum (Hvam & Toft
which may last several months, males remain most of the time 2008; Table 4). There is a parallel to this in bird trophic
inside their mud nests protecting their eggs and waiting for the biology. Experiments in which certain bird species were
visit of ovigerous females (Mora 1990; Quesada-Hidalgo et al. offered only fruit diets, likewise have shown that individuals
2019). Although paternal care constrains foraging activity of lost weight, and this was attributed to the extreme lipid and
the caring males, their body condition does not decrease over protein deficiency of a low-quality fruit diet (Bairlein 1996;
time (Requena & Machado 2015). This finding contrasts with Jordano 2000).
another harvestman species with exclusive paternal care, For spiders, no experiment like the one reported above for
Iporangaia pustulosa Mello-Leitão, 1935 (Gonyleptidae), in harvestmen has been performed so far. However, it can be said
which the body condition of caring males deteriorates over that, contrary to harvestmen, consumption of fruit pulp and
time (Requena et al. 2012). One possible explanation for this seed appendages by spiders under natural conditions is
difference is that caring males of Quindina regularly feed on apparently uncommon, so that this issue seems to be of little
the fungi that grow inside and around their nests (Figs. 2C–D) relevance to this arachnid group. Bagheera kiplingi Peckham &
whereas caring males of I. pustulosa most likely do not have Peckham, 1896 (Salticidae) is the only spider species known so
access to this food source (Requena & Machado 2015). In the far to exhibit a predominantly vegetarian lifestyle (Meehan et
future, it could be tested whether caring males of Quindina that al. 2009). But even this species, which feeds to a large extent on
are somehow prevented from feeding on fungi are able to the detachable leaf tips of Acacia trees (i.e., Beltian bodies)
sustain their body condition over the period of parental care. and nectar, individuals perish after one to several weeks and
We are not aware of any experimental study in which always before molting to the next instar if kept strictly on a
harvestmen were fed with exclusively mushrooms and we plant-based diet (Nyffeler et al. 2016).
therefore do not know how they would respond under such
unbalanced nutritional conditions. For spiders, this question 5. CONCLUDING REMARKS
has been tested by means of laboratory feeding trials.
Spiderlings fed a diet consisting of only fungal spores Like scorpions, harvestmen are ‘‘solid food feeders’’, which
(Cladosporium sp.) did not live longer than starving spider- means that they can consume solid tissue of mushrooms, fruit
lings, suggesting that the spores were of no nutritional value. pulp, seeds, or seed appendages as a food source by biting off
Similar experimental findings have been reported for other small pieces. Another possibility could be that harvestman
arthropods. If a diet consisting of only fungal spores would need alternative sources of food as they are typically
(Cladosporium sp. and two other fungal species) was offered bad at capturing prey compared to other arachnids. The fact
to honeybees, this type of food was rejected (Parish et al. that 3.4 times as many records of harvestmen feeding on
2020). In another study, consumption of fungal spores by mushrooms and fruits could be found as compared to the
honeybees was shown to be detrimental, suggesting that the spiders (96 vs. 28 records; Tables 1–3) appears to signify this
spores contain some type of toxic secondary compound difference in the foraging behavior of the two arachnid groups.
(Schmidt et al. 1987). Fungal spores appear to be low-quality Although no experimental studies on harvestmen have been
food probably because they have rigid protective walls – made conducted to test whether individuals can survive on an
up of complex three-dimensional network of polysaccharides – exclusively vegetarian diet, we anticipate that the answer is
that make it difficult to access the nutritious inner content probably no, because fungi, fruit pulp, seeds, and seed
(Lundgren 2009; Noothalapati et al. 2016). In addition, fungal appendages may lack some micro- and macro-nutrients, as
spores have low contents of protein and moisture compared to well as vitamins, that are exclusively acquired from animal
mushrooms. Thus, fungivory appears to be of little relevance prey (Simpson & Raubenheimer 2012). However, mushrooms,
to spiders from a nutritional point of view (see subsection ‘‘4.3 fruit pulp, seeds, and/or seed appendages can make up a
Nutritional benefits of fungivory’’ above). substantial portion of the natural diet of harvestmen at times
Regarding frugivory, studies conducted with Neotropical when these food sources are available in high abundance
harvestmen from the families Cosmetidae and Gonyleptidae (Machado & Pizo 2000; Schaus et al. 2013; Waldenmaier
suggest that opportunistic frugivory may be important 2016). Detailed information on the diet of a larger number of
species and families will certainly show that consumption of on an elaiosome. We also wish to thank Luis E. Acosta
vegetarian food items is widespread in the order Opiliones. (Universidad Nacional de Córdoba, Argentina) for helping to
In the case of spiders, fungivory and frugivory appear to be get access to literature and Norberto Palacios for helping to
of little relevance for their diet. This might be partially scan some old slides. Valuable comments of the subject editor
explained by the fact that spiders are ‘‘fluid feeders’’ (Foelix Yael Lubin (Ben-Gurion University) and two anonymous
2011). In fact, spiders feed on vegetarian diets most frequently reviewers helped to improve the manuscript. Last but not
in the form of fluids, such as nectar, stigmatic exudate, plant least, we thank the photographers Susannah Anderson
sap, and honey dew (e.g., Pollard et al. 1995; Taylor & Foster (Canada), Gilbert Barrantes (Costa Rica), Nicky Bay (Singa-
1996; Jackson et al. 2001; Nyffeler et al. 2016; Suzuki & Sano pore), Isa S. Betancourt (USA), Rhett Butler (Mongabay,
2021). Solid mushroom and fruit tissues are, therefore, not USA), Bruno A. Buzatto (Brazil), Hock Ping Guek (Malay-
suitable food sources for spiders. An exception occurs when sia), Ryosuke Kuwahara (Japan), Sean McCann (USA),
synanthropic spiders feed on fruit waste, because in this Marco Aurélio Pizo (Brazil), Gustavo S. Requena (Brazil),
particular situation the juice of fruits cut open by human Andrés Rojas (Costa Rica), Chuck Rosenberger (USA), Bruce
residents can easily be accessed by the spiders. How frequently Rottink (USA), Martin Suvák (Slovakia), John A. Uribe
spiders feed on the liquefied state of the flesh of fruits in (Brazil), Rodrigo H. Willemart (Brazil), and Melvyn Yeo
natural systems is still unexplored. However, we anticipate (Singapore).
that fruits probably do not represent a relevant food item for
species of the order Araneae. SUPPLEMENTAL MATERIALS
ACKNOWLEDGMENTS Available online at https://doi.org/10.1636/JoA-S-22-015.s1

Appendix 1.—Combinations of key words used in the
North American harvestmen were identified by James C. literature search in all databases.
Cokendolpher (Texas Tech University, USA) and Jeffrey W. Figure S1.— Further cases of fruit consumption by
Shultz (University of Maryland, USA). Harvestmen from harvestmen.
Brazil, Costa Rica, Malaysia, and Singapore were identified by
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Journal of Arachnology 51:1–18

Fungus and fruit consumption by harvestmen and spiders (Opiliones, Araneae):

1. INTRODUCTION Species in these orders exhibit an enormous diversity of lifestyles

Arachnid taxon Plant species Plant family Country No of records Reference

Arachnid taxon Plant species Plant family Country No of records Reference

– Galetti et al. (2000): Fruit data for Euphorbiaceae, 3. RESULTS

Arachnid Water Lipids Protein Carbohydrates kcal/100 Avg. fruit weight

ACKNOWLEDGMENTS Available online at https://doi.org/10.1636/JoA-S-22-015.s1

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