Flying squirrels use a mortise-tenon structure to fix nuts on understory twigs

  1. Han Xu
  2. Lian Xia
  3. John R Spence
  4. Mingxian Lin
  5. Chunyang Lu
  6. Yanpeng Li
  7. Jie Chen
  8. Tushou Luo
  9. Yide Li
  10. Suqin Fang  Is a corresponding author
  1. Research Institute of Tropical Forestry, Chinese Academy of Forestry, China
  2. College of Forestry, Hainan University, China
  3. Department of Renewable Resources, University of Alberta, Canada
  4. State Key Laboratory of Biocontrol, School of Life Sciences, Sun Yat-Sen University, China
11 figures, 15 videos, 3 tables and 2 additional files

Figures

Spatial distribution of the 151 suspended nuts observed in Jianfengling Nature Reserve, Hainan, China.
Nuts are stored after surface preparation by flying squirrels.

(A) Nut of C. edithiae (Skan) Schottky, with chewed grooves outlined in red. Nuts of C. edithiae fixed on trees, with (B–D) one groove, (E) two non-connected grooves, or (F) spiral carved grooves encircling the nuts. (G) Nut of C. patelliformis (Chun) Y. C. Hsu et H. W. Jen, with chewed grooves outlined in red. (H–I). Nuts of C. patelliformis fixed on trees, with carved grooves on the bottom fixed on (J) bamboos, (K–L) lianas, between the big petioles of (M) trees and (N) palms.

Variation in carved grooves depends on the storage situation.

The carved surface grooves on nuts of C. edithiae mostly encircle the middle of the nut, with (A) one groove, (B) one spiral groove, or (C) two separated grooves. The grooves on nuts of C. patelliformis are distributed on the bottom of the nuts, with (D) 2, (E) 4, (F) 6, (G) 8, (H) 10 symmetrically, or (I) randomly distributed grooves.

Nuts were fixed tightly between twigs generally meeting at angles of 25–40°.

(A) C. edithiae nuts. (B) C. patelliformis nuts.

Most nuts were stored on small plants with the diameter at breast height (DBH) ranging from 0.4–1.6 cm.

(A) C. edithiae nuts. (B) C. patelliformis nuts. Notes: The value on each bar is the actual number of stored nuts.

Nuts were stored mainly on small plants between twigs with diameters of 0.10 - 0.60 cm.

(A) Histogram of diameters of twigs used to store nuts of C. edithiae. (B) Histogram of diameters of twigs used to store nuts of C. patelliformis. Notes: The value on each bar is the actual twigs with the number of stored nuts.

Grooves carved by squirrels on most C. edithiae nuts were 0.2 - 0.6 cm in width.
Nuts were generally stored on the first to third branches at 1.5–2.5 m aboveground.

(A) C. edithiae nuts. (B) C. patelliformis nuts. Notes: The value on each bar is the actual number of stored nuts.

Number of grooves carved on the oblate nuts of C. patelliformis.

(A) Most nuts had fewer than eight grooves. (B) The depth of most grooves was shallow to medium. Notes: The value on each bar is the actual number of stored nuts.

Distance from storage sites to potential parent trees for the nuts varied from 10–25 m.

(A) C. edithiae nuts. (B) C. patelliformis nuts. Notes: The value on each bar is the actual number of stored nuts.

After long (e.g. >ca. 365 days) storage, nuts become not fresh.

(A) Dried nuts, (B) Germinated, or (C–E) Destroyed by insects.

Videos

Video 1
Squirrel 1 of Hylopetes alboniger was checking and re-fixing nuts at the storage sites with footage from infrared cameras.
Video 2
Squirrel 2 of Hylopetes alboniger was removing nuts from storage sites with footage from infrared cameras.
Video 3
Squirrel 3 of Hylopetes alboniger was removing nuts from storage sites with footage from infrared cameras.
Video 4
Squirrel 4 of Hylopetes phayrei electili was checking and re-fixing nuts at storage sites with footage from infrared cameras.
Video 5
Squirrel 5 of Hylopetes phayrei electili was removing nuts from storage sites with footage from infrared cameras.
Video 6
One squirrel was cracking the nuts on the ground with footage from infrared cameras.
Video 7
One squirrel was rotating and cracking the nuts on the trees with footage from infrared cameras.
Video 8
One squirrel was re-fixing the nuts by cracking behavior with footage from infrared cameras.
Video 9
One squirrel was fixing the nuts between the twigs with footage from infrared cameras.

We merged several photos and a video successively taken by an infrared camera in 30 s.

Video 10
Footage of shaking a liana does not dislodge nuts of Cyclobalanopsis edithiae stored by squirrels.
Video 11
Footage of shaking a sapling does not dislodge nuts of Cyclobalanopsis edithiae stored by squirrels.
Video 12
Footage of shaking a sapling does not dislodge nuts of Cyclobalanopsis edithiae stored by squirrels.
Video 13
Footage of shaking a liana does not dislodge nuts of Cyclobalanopsis patelliformis stored by squirrels.
Video 14
Footage of shaking a sapling does not dislodge nuts of Cyclobalanopsis patelliformis stored by squirrels.
Video 15
Footage of shaking a sapling does not dislodge nuts of Cyclobalanopsis patelliformis stored by squirrels.

Tables

Table 1
The nine squirrel species known from Jianfengling, Hainan Island, China.
Species and subspecies nameBody length/mm
Tamiops maritimus (Bonhote,1900) (Liu et al., 2020; Pan et al., 2007)105~134
Dremomys pyrrhomerus (Thomas, 1895) (Xu and Chen, 1989)194~215
Hylopetes alboniger (Hodgson, 1870) (Liu et al., 2020; Andrew, 2008; Pan et al., 2007)180~203
Hylopetes phayrei electilis (Allen, 1925) (Liu et al., 2020; Pan et al., 2007)123~173
Dremomys rufigenis (Blanford, 1878) (Zheng et al., 2008)170~250
Callosciurus erythraeus (Pallas, 1779) (Huang, 1995; Zheng et al., 2008)198~252
Belomys pearsonii (Gray, 1842) (Pan et al., 2007; Huang, 1995)180~260
Petaurista albiventer (Gray, 1834) (Jing et al., 2007)420~520
Ratufa bicolor (Sparrmann, 1778) (Li et al., 2008)350~505
  1. Note: The data in this table are referenced from the below literature.

Table 2
The types of plants used for nut storage.
Plant typeNumber of individualsPercentage of all individuals (%)
Alive tree10871.5
Dead tree1711.3
Alive liana1912.6
Dead liana21.3
Bamboo53.3
Total151100
Table 3
Main Fagaceae species found in a 60 ha plot in the Jianfengling forest.
SpeciesAbundance
Castanopsis carlesii (Hemsley) Hayata3269
Castanopsis fissa (Champion ex Bentham) Rehder & E. H. Wilson2803
Castanopsis jianfenglingensis Duanmu2297
Castanopsis tonkinensis Seemen953
Castanopsis ledongensis C. C. Huang & Y. T. Chang335
Castanopsis fabri Hance113
Castanopsis hystrix J. D. Hooker & Thomson ex A. de Candolle35
Cyclobalanopsis edithiae (Skan) Schottky1645
Cyclobalanopsis patelliformis (Chun) Y. C. Hsu & H. W. Jen1207
Cyclobalanopsis phanera (Chun) Y. C. Hsu & H. W. Jen886
Cyclobalanopsis fleuryi (Hickel & A. Camus) Chun ex Q. F. Zheng568
Cyclobalanopsis neglecta Schottky392
Cyclobalanopsis blakei (Skan) Schottky279
Cyclobalanopsis hui (Chun) Chun ex Y. C. Hsu & H. W. Jen220
Lithocarpus longipedicellatus (Hickel & A. Camus) A. Camus2842
Lithocarpus pseudovestitus A. Camus2427
Lithocarpus fenzelianus A. Camus1751
Lithocarpus amygdalifolius (Skan) Hayata1360
Lithocarpus handelianus A. Camus1046
Lithocarpus fenestratus (Roxburgh) Rehder323
Lithocarpus howii Chun130
Lithocarpus hancei (Benth.) Rehd.71

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  1. Han Xu
  2. Lian Xia
  3. John R Spence
  4. Mingxian Lin
  5. Chunyang Lu
  6. Yanpeng Li
  7. Jie Chen
  8. Tushou Luo
  9. Yide Li
  10. Suqin Fang
(2023)
Flying squirrels use a mortise-tenon structure to fix nuts on understory twigs
eLife 12:e84967.
https://doi.org/10.7554/eLife.84967